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ILLINOIS  BIOLOGICAL 
MONOGRAPHS 


PUBLISHED  QUARTERLY 

UNDER  THE  AUSPICES  OF  THE  GRADUATE  SCHOOL 

BY  THE  UNIVERSITY  OF  ILLINOIS 


VOLUME  I 


Urbana,  Illinois 
1914-15 


Editorial  Committee 


Stephen  Alfred  Forbes  William  Trelease 

Henry  Baldwin  Ward 


\ 


TABLE  OF  CONTENTS 

Volume  I 


NUMBERS  PAGES 

i  and  2    A  Revision  of  the  Cestode  Family  Proteocephalidae.    By  George 

Roger  La  Rue.    With  16  plates i-35<> 


( Distributed  November  30,  1914) 


3        Studies    on    the   Cestode   Family   Anoplocephalidae.     By    Herman 

Douthitt.    With  6  plates 351-446 

(Distributed  March  8,  1915) 


4        Some  North  American  Larval   Trematodes.     By   William  Walter 

Cort.    With  8  plates 447-532 

(Distributed  June  28.  1915) 


732170 


ILLINOIS  BIOLOGICAL 
MONOGRAPHS 


Vol.1  January,  1915  '  No.  3 


Editorial  Committee 


Stephen  Alfred  Forbes  William  Trelease 

Henry  Baldwin  Ward 


Published  under  the 

Auspices  of  the  Graduate  School  by 

the  University  of  Illinois 


Copyright,  1915 
By  the  University  of  Illinois 


STUDIES  ON 

THE    CESTODE    FAMILY 

ANOPLOCEPHALID^ 


WITH  SIX  PLATES 


BY 


HERMAN  DOUTHITT 


Contributions  from  the 

Zoological   Laboratory  of  the  University   of  Illinois  under  the  direction  of 

Henry   B.  Ward,  No.  38 


TABLE  OF  CONTENTS 

PAGE 

Introduction    5 

Anatomical  Discussions  of  Species  and  Genera 

Andrya  primordialis 5 

Andrya  communis 8 

Andrya    macrocephala 10 

Andrya  translucida 13 

The  Genus  Andrya 16 

Anoplocephala  wimerosa 17 

Anoplocephala   variabilis 20 

Anoplocephala  infrequens 24 

The  Genus  Anoplocephala 26 

Schizotaenia   americana 28 

Schizotaenia  variabilis .• _...  33 

Schizotaenia  anoplocephaloides 35 

The  Genus  Schizotaenia 40 

The  Genus  Moniezia,  and  the  uterus  of  Moniezia  expansa 43 

The  Cittotaeniae  of  North  American  Rabbits 46 

The  Genus  Cittotaenia 49 

Comparative  Studies  on  the  Anoplocephalidae 

Comparative  Anatomy  of  the  Anoplocephalinae SO 

Evolution  of  the  Anoplocephalidae 54 

Observations  on  the  Life  Histories  of  Anoplocephaline  Parasites  of  the  Pocket 

Gopher .' 60 

Key  to  Known  Species  of  the  Anoplocephalidae 63 

Summary  and  Conclusions 79 

Bibliography 81 

Explanation  of  Plates „ 85 


355]  ANOPLOCEPHALIDAE— DOUTHITT 


INTRODUCTION 

The  present  paper  gives  the  results  of  studies  begun  at  the  Univer- 
sity of  Illinois  in  December,  1910,  and  carried  on,  between  numerous 
interruptions,  to  the  present  time. 

My  purpose  has  been  to  make  a  comparative  anatomical  study  of 
the  Anoplocephalidae,  but  since  so  few  of  the  individual  representatives 
of  the  family  have  received  adequate  study,  it  has  been  necessary  for  me 
to  give  most  of  the  time  to  the  individual  study  of  undescribed  or  poorly 
described  species.  The  work  lacks  much  of  being  complete;  many  of 
the  genera  I  have  not  had  opportunity  to  study  myself.  Other  work 
compelled  me  to  lay  the  task  aside,  and  it  seems  advisable  to  make  public 
the  results  already  obtained. 

My  thanks  are  due  to  Professor  Henry  B.  Ward  for  assistance  of 
many  sorts;  to  Professors  Robert  T.  Young,  M.  J.  Elrod,  and  R.  A. 
Lyman,  Drs.  B.  H.  Ransom,  and  John  E.  Gutberlet,  for  materials  placed 
at  my  disposal  for  study;  also  to  Miss  Bertha  E.  Martin  for  criticisms 
and  assistance  in  preparing  manuscript. 


ANATOMICAL  DESCRIPTION  OF  SPECIES  AND  GENERA 
Andrya  primordialis  sp.  nov. 
[Figures  1-4] 

Two  specimens  of  this  cestode,  one  without  scolex,  were  taken  from 
a  red  squirrel  (Sciurus  hudsonica)  at  Bemidji,  Minnesota,  in  Septem- 
ber, 1911.  Since  about  20  squirrels  in  all  were  examined,  there  and  at 
Brainerd,  Minnesota,  it  seems  that  this  species  is  rare,  at  least  as  far  as 
this  region  and  host  are  concerned.  Other  species  of  squirrels  from 
these  and  other  localities  were  likewise  uninfected  with  this  species. 
Its  close  relationship  to  the  ancestral  types  of  the  Anoplocephalidae 
seems  evident,  even  tho  based  upon  a  study  of  so  few  specimens. 

The  worms,  neither  of  which  are  fully  grown,  have  a  length  of  40 
mm.  and  the  single  complete  specimen  has  155  proglottids.  The  strobila 
increases  in  width  to  the  posterior  end,  there  being  1  mm.  in  breadth. 
The  first  proglottids  are  290^,  wide  and  one-eighth  as  long.  Mature 
proglottids  are  half  as  long  as  wide,  and  the  proglottids  farther  back 


6  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [356 

are  somewhat  longer.  The  scolex  is  compact,  700/u,  long  by  475/x  broad, 
the  greatest  width  being  near  the  anterior  end.  Posteriorly  the  scolex 
tapers  uniformly,  until  it  merges  into  the  neck,  which  is  290/x  broad. 
The  suckers  open  nearly  directly  forwards.  A  circular  groove  about 
the  scolex  at  the  level  of  the  suckers  makes  it  appear  as  if  the  tip  of 
the  scolex  were  contracted  somewhat  within  the  rest. 

The  genital  pore  is  always  on  the  right  margin,  about  four-fifths 
the  length  of  the  proglottid  from  its  anterior  end.  It  is  relatively  large 
and  deep  as  compared  with  the  same  organ  in  related  cestodes.  The 
cirrus  pouch  is  much  larger  than  in  other  known  members  of  the  genus, 
both  in  length  and  breadth,  its  median  end  lying  well  across  the  excre- 
tory ducts.  Its  inner  end  is  rounded  and  its  lateral  end  only  slightly 
tapering.  Across  the  middle  its  diameter  is  considerably  less  than  at 
either  end  so  that  its  outline  is  somewhat  like  that  of  the  figure  8.  The 
median  end  is  occupied  by  the  vesicula  seminalis.  The  vas  deferens  is 
not  enlarged  nor  greatly  coiled.  An  elongated  prostate  gland  lying  in 
the  anterior  end  of  the  proglottid  empties  into  the  vas  deferens  near 
the  cirrus  pouch.  The  testes  are  dorsal,  occupying  the  entire  median 
field  to  the  left  of  and  anterior  to  the  vitelline  gland  and  receptaculum 
seminis,  and  partly  underlying  the  latter.  In  the  distal  half  of  the 
left  side  the  testicular  field  extends  laterad  beyond  the  ventral  excre- 
tory duct.  The  testes  number  30  to  40  and  are  mostly  70  to  80/x  long, 
the  breadth  being  somewhat  less.  They  break  down  early,  but  the 
membranes  remain  intact  and  the  individual  testes  are  easily  distin- 
guishable. 

The  vagina  lies  entirely  posterior  to  the  cirrus  pouch,  and  in  the 
same  dorsoventral  plane.  Its  coils  are  thickly  beset  with  glandular 
cells.  The  receptaculum  seminis  is  large  and  slightly  longer  anteropos- 
teriorly  than  laterally.  It  lies  directly  in  front  of  the  vitelline  gland, 
and  does  not  reach  laterad  to  the  ventral  excretory  duct  as  is  the  case 
in  other  species  of  the  genus.  The  ovary  is  a  semicircular  mass,  reaching 
nearly  across  the  median  field.  In  immature  stages  12  to  15  lobes  can 
be  distinguished  which  radiate  in  all  horizontal  directions;  but  before 
the  ovary  matures  these  lobes  become  indistinguishable.  The  ovary  is 
not  located  nearer  the  pore  side  than  the  other,  the  mouth  of  the  oviduct 
being  as  often  to  the  left  as  to  the  right  of  the  median  line.  The 
vitelline  gland  however  is  distinctly  nearer  the  pore  side.  It  is  of  the 
shape  most  usually  found  in  this  group,  being  composed  of  a  large 
median  and  a  smaller  lateral  portion  connected  by  a  transverse  portion. 

The  uterus,  just  before  sexual  maturity,  is  a  network  of  tubes  in 
the  anterior  two-thirds  of  the  proglottid.  In  the  median  field  it  extends 
distad  to  below  the  upper  limit  of  the  ovary,  ventrad  of  the  latter.    The 


357]  ANOPLOCEPHALIDAi—DOUTHITT  7 

lateral  tips  extend  a  little  farther  distad  than  the  median  portion  and 
cross  the  excretory  ducts  ventrally  on  either  side.  The  anterior  and 
lateral  margins  together  form  a  semicircle,  so  that  at  this  stage  the 
uterus  as  a  whole  is  somewhat  semilunar  in  outline.  The  early  develop- 
ment of  the  uterus  will  be  taken  up  later  (page  51).  Development 
from  the  stage  just  described  is  first  by  enlargement  and  coalescence  of 
tubes,  forming  a  saccular  structure ;  then  by  regular  outpocketing,  ante- 
riorly, distally,  and  laterally.  The  anterior  and  lateral  pockets  together 
number  about  22;  the  distal,  15.  Since  neither  specimen  possessed 
completely  ripe  proglottids,  a  study  of  the  embryo  was  unfortunately 
not  possible. 

The  dorsal  excretory  duct  lies  laterad  of  the  ventral  and  usually 
somewhat  more  dorsal.  The  diameter  of  the  ventral  is  30/*  and  that  of 
the  dorsal  7/x.  The  transverse  commissure  is  of  very  small  diameter 
and  is  not  visible  at  all  in  sections  at  hand  except  in  proglottids  past 
sexual  maturity. 

The  presence  of  the  prostate  gland  and  the  reticulate  uterus  show 
at  once  that  this  species  is  allied  to  the  genus  Andrya.  It  disagrees 
however  with  the  accepted  diagnosis  in  that  (1)  the  pores  are  strictly 
dextral,  (2)  the  ovary  is  not  nearer  the  pore  side  than  the  opposite  side, 
(3)  the  testes  extend  laterad  across  the  ventral  excretory  duct  on  the 
aporose  side;  and  (4)  the  later  development  of  the  uterus  is  by  regular 
anterior  and  posterior  outpocketing. 

The  first  point  of  disagreement  is  not  an  important  one  since 
unilaterality  is  approached  in  all  of  the  species  of  the  genus.  The  sec- 
ond point,  while  significant  in  other  aspects  cannot  be  regarded  as  of 
generic  import  since  the  other  female  glands  are  displaced  in  the  regular 
manner.  The  third  point  will  be  shown  to  be  true  also  of  known  and 
new  species  of  Andrya,  Anoplocephala,  and  Bertiella;  so  that  the 
descriptions  of  these  genera  are  in  error,  and  this  character,  supposed 
to  be  confined  to  the  genus  Aporina,  is  found  in  all  the  single-pored 
genera  of  the  subfamily  except  Schizotaenia.  As  to  the  fourth,  the 
discrepancy  seems  to  be  due  to  errors  in  the  descriptions  of  the  species 
already  known;  for  while  Stiles  (1896),  describing  the  uterus  of  Andrya 
rhopalocephala  says  it  is  in  its  final  stages  a  simple  sac  or  with  ' '  at  most 
extremely  fragmentary  and  rudimentary  divisions",  his  figures  of 
gravid  uteri  of  this  species  and  A.  cuniculi  (see  his  Plate  VIII,  fig.  1 
and  Plate  IX,  fig.  1)  show  for  both  species  what  is  apparently  regular 
outpocketing,  correctly  drawn  tho  misinterpreted. 

This  cestode  shows  several  characters  which  mark  it  as  the  most 
primitive  species  known  in  the  genus;  and  if  the  contention  later  made 


8  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [358 

be  accepted,  that  Andrya  is  the  most  primitive  genus  of  the  family,  then 
this  becomes  the  most  primitive  known  species.  Its  primitive  characters 
are  (1)  the  very  broad  distribution  of  the  testes  which  occupy  nearly 
the  entire  median  field  and  part  of  the  lateral;  (2)  the  fact  that  the 
ovary  is  not  nearer  the  pore  side  than  the  opposite  side;  and  (3)  the 
limitation  of  the  receptaculum  seminis  to  the  median  field  and  its  simple 
globular  character.  In  these  characters  this  species  is  approached  in 
some  respects  by  A.  communis,  next  to  be  described,  which  must  be 
regarded  as  its  nearest  known  ally. 


Andrya  communis  sp.  nov. 
[Figures  5-8] 

Professor  R.  T.  Young  collected  in  1908  at  Long's  Peak,  Colorado, 
cestodes  from  Evotomys  gapperi  galei,  Microtus  pennsylvanicus  modes- 
tus  and  Peromyscus  sp.  They  were  examined  by  Hall  who  concluded 
and  reported  (1912)  that  those  from  Evotomys  and  Microtus  repre- 
sented a  species  of  Anoplocephala,  while  those  from  Peromyscus  rep- 
resented a  second  species  of  the  same  genus.  Professor  Ward  has  secured 
for  me  the  loan  of  this  material  for  study.  The  cestodes  from  Pero- 
myscus belong  really  in  the  genus  Hymenolepis,  except  for  one  slide, 
apparently  mislabelled,  which  is  of  the  same  species  as  those  from 
Microtus.  Those  from  Evotomys  and  Microtus  prove  to  be  two  distinct 
species,  the  first  being  an  Andrya  and  the  other  probably  an  Andrya 
also,  tho  the  fragments  at  hand  are  all  too  far  past  sexual  maturity 
to  allow  of  certain  generic  determination.  The  latter  species  must 
therefore  of  necessity  be  omitted  from  consideration,  tho  it  comes  within 
the  scope  of  this  paper.     . 

The  species  under  consideration  from  Evotomys  gapperi  galei  is 
represented  in  the  material  at  hand  by  about  200  fragments  in  alcohol 
and  17  slides,  most  of  which  were  prepared  by  Professor  Young.  Only 
one  scolex  is  in  this  lot,  this  being  in  the  form  of  oblique  cross  sections. 
Recently  I  have  found  in  the  collection  of  Professor  Ward  four  frag- 
ments, evidently  representing  one  worm,  which  were  of  the  same  lot  of 
material,  an. I  were  presented  to  Professor  Ward  several  years  ago  by 
Professor  Young.  This  material  includes  a  scolex  with  a  considerable 
number  of  proglottids  attached,  thus  furnishing  an  idea  of  the  appear- 
ance of  the  complete  worm. 

The  total  length  is  estimated  to  be  about  3  or  4  cm;  the  number  of 
proglottids,  about  225.  The  greatest  width  is  about  1.5  to  2  mm.  The 
proglottids  vary  in  shape  from  12  times  as  long  as  broad  shortly  before 


359]  ANOPLOCEPHALIDAZ—DOUTHITT  9 

sexual  maturity  to  longer  than  broad  in  the  case  of  ripe  proglottids. 
The  length  of  the  scolex  is  roughly  560/x.  The  greatest  width,  350/*,  is 
reached  95/a  from  the  anterior  end.  The  anterior  end  is  rounded; 
posteriorly  the  scolex  narrows  regularly. 

The  genital  pores  are  all  on  the  right  margin  near  the  middle  of 
the  proglottid.  The  circular  muscles  surrounding  the  inner  end  of  the 
cirrus  pouch  are  usually  contracted,  compressing  this  part  and  making 
the  pouch  inversely  pear-shaped.  In  older  proglottids  however  the 
inner  end  becomes  distended  and  much  larger  than  the  outer.  In  some 
specimens  the  cirrus  pouch  is  separated  by  a  distance  equal  to  its  own 
length  from  the  ventral  excretory  duct ;  in  others  its  median  end  extends 
across  both  ducts  (Figs.  5  and  8).  The  different  conditions  do  not 
seem  to  depend  so  much  on  the  stage  of  contraction  of  the  proglottid 
as  upon  differences  in  the  position  of  the  excretory  ducts.  An  internal 
vesicula  seminalis  is  present.  The  cirrus  is  generally  extruded,  tho 
usually  it  does  not  extend  outside  the  genital  pore. 

The  vas  deferens  is  not  enlarged  and  is  not  coiled,  tho  it  is  probable 
that  some  coiling  would  be  found  in  more  expanded  specimens.  A 
distinct  prostate  gland  is  present ;  it  is  tubular  with  a  globular  enlarge- 
ment at  its  end.  Lateral  to  the  pore  of  the  prostate  gland  the  vas 
deferens  is  surrounded  by  a  loose  cellular  mesh.  The  testes  are  all 
dorsal.  On  the  left  they  may  extend  across  both  excretory  ducts,  or 
they  may  be  strictly  limited  to  the  median  field.  To  the  right  they 
extend  as  far  as  the  prostate  gland  in  the  anterior  part  of  the  field. 
They  extend  distad  as  far  as  the  vitelline  gland  and  receptaculum 
seminis,  and  on  the  left  of  these  they  extend  to  the  distal  end  of  the 
proglottid.  The  number  varies  in  eight  proglottids  counted  from  24 
to  41.  They  are  mostly  tranversely  elongated,  55  to  80 fi  long  in  trans- 
verse axis  by  30  to  50/x  in  longitudinal  axis. 

The  vagina  lies  entirely  posterior  to  the  cirrus  pouch  and  vas 
deferens  and  in  the  same  plane.  Up  to  the  beginning  of  the  receptacu- 
lum seminis  its  walls  are  glandular.  The  receptaculum  begins  where 
the  vagina  crosses  the  dorsal  excretory  duct;  it  enlarges  gradually  and 
uniformly  to  near  its  inner  end  then  becomes  abruptly  smaller.  The 
ovary  is  an  asymmetrical  mass  most  of  whose  bulk  lies  on  the  pore  side, 
but  it  approaches  no  nearer  to  the  excretory  ducts  on  one  side  than  on 
the  other  and  the  oocapt  is  as  often  to  the  left  as  to  the  right  of  the 
median  line.  It  shows  rather  obscure  and  uncertain  evidence  of  partial 
division  into  about  seven  radiating  lobes.  The  lateral  and  median  lobes 
of  the  vitelline  gland  are  but  little  larger  than  the  transverse  portion 
of  the  gland  which  connects  them.  This  gland  lies  considerably  to  the 
pore  side  of  the  median  line. 


10  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [360 

The  uterus  is  a  typical  reticulum  in  its  early  stages.    It  lies  entirely 
anterior  to  and  to  the  left  of  the  ovary,  being  never  ventrad  of  it.    To 
the  left  of  the  ovary  it  reaches  the  transverse  commissure  posteriorly. 
On  either  side  it  crosses  the  ventral  excretory  ducts  ventrally.    Devel 
opment  from  this  stage  is  as  in  A.  primordialis. 

The  excretory  ducts  are  dorsal  and  ventral  in  position  in  the  head 
region;  but  the  dorsal  duct  very  early  moves  to  a  position  laterad  of 
the  ventral,  which  it  retains.  The  ventral  duct  is  four  times  the  diame- 
ter of  the  dorsal ;  the  transverse  commissure  is  very  small.  The  ventral 
duct  diverges  from  the  central  axis  as  it  passes  distad  until  near  the 
distal  end  of  the  proglottid  where  it  turns  almost  directly  mediad  to 
regain  its  position.  The  dorsal  duct  has  a  somewhat  similar  course  but 
not  so  pronounced. 

This  cestode  it  will  be  seen  is  rather  closely  related  to  Andrya 
primordialis.  Many  of  the  differences  in  the  two  accounts  are  due 
probably  to  differences  in  the  state  of  contraction  in  the  material. 

Andrya  macrocephala  sp.  nov. 
[Figures  9-13] 

Fifteen  specimens  of  this  species  were  taken  from  pocket  gophers 
(Geomys  bursarius)  living  in  swampy  river  bottom  land  at  Brainerd, 
Minnesota,  and  one  from  a  gopher  living  in  low,  heavy,  black  soil  at 
Thief  River  Falls,  Minnesota.  Apparently  they  are  entirely  absent 
from  sandhill  regions  if  not  from  all  uplands.  For  distribution  and 
frequency  see  the  table  on  page  62. 

By  far  the  most  conspicuous  anatomical  feature  of  this  cestode  is 
the  enormous  development  of  the  ventral  excretory  ducts  and  commis- 
sures. In  the  scolex  anterior  to  the  distal  end  of  the  suckers,  the  four 
ducts  have  a  winding  course,  the  diameter  of  each  varying  generally 
between  18  and  32/x.  Distad  of  the  suckers  the  ducts  have  a  straight 
course,  are  approximately  15/t  in  diameter,  and  are  arranged  as  a  dorsal 
and  a  ventral  pair.  Near  the  distal  end  of  the  scolex  the  dorsal  duct 
begins  to  move  laterad  and  soon  comes  to  lie  directly  laterad  of  the 
ventral.  The  diameter  of  the  dorsal  duct  throughout  the  strobila  is 
about  18/i.  The  ventral  ducts  increase  considerably  in  size  as  they  pass 
distad.  As  the  proglottids  approach  sexual  maturity  this  increase  be- 
comes much  more  rapid  and  the  transverse  commissures  appear  and 
develop  rapidly.  In  a  sexually  mature  proglottid  the  ventral  aporose 
duct  near  the  middle  of  the  proglottid  is  80/i  in  transverse  diameter; 
at  either  end  it  is  145/a  in  diameter.  On  the  pore  side  the  transverse 
diameter  is  125/t  in   the  center;  at  either  end,  150/t  or  more.     The 


361]  ANOPLOCEPHALIDJE—DOUTHITT  11 

dorsoventral  diameter  in  the  center  is  225/t.  The  transverse  commissure 
measures  at  its  largest  point  about  100/x;  at  either  end  it  becomes 
smaller.  As  the  proglottids  age  the  ducts  keep  on  increasing  in  size. 
The  largest  dimensions  observed  for  the  transverse  commissure  were 
320/a;  for  longitudinal  ducts,  exclusive  of  interproglottidal  enlarge- 
ments, both  transverse  and  dorsoventral  diameters  reached  320//,;  for 
interproglottidal  enlargements,  430/x.  As  the  proglottids  lengthen  prior 
to  detachment,  the  ducts  become  smaller  as  the  result  of  the  stretching, 
and  the  interproglottidal  enlargements  which  are  not  thus  affected  be- 
come prominent  as  spindle-shaped  objects. 

Since  the  maximum  breadth  of  this  worm  is  1.5  mm.  it  will  be  seen 
-that  this  development  is  truly  enormous,  the  two  ventral  ducts  extend- 
ing in  some  cases  through  not  far  from  one  half  the  transverse  measure- 
ment of  the  proglottid  and  dorsoventrally  through  the  cortical  layer. 
This  is  not  due  to  reagents  since  the  living  worms  exhibit  the  enormous 
ducts  clearly  visible  to  the  naked  eye,  before  being  given  any  treatment 
whatever.  Great  as  this  development  is  however  it  will  be  seen  that  it 
is  exceeded  in  Andrya  translucida,  the  next  form  to  be  considered,  tho 
with  the  difference  that  in  the  present  species  the  dimensions  are  con- 
stant in  any  given  stage  of  development  of  the  proglottid  while  in  the 
other  they  will  suddenly  develop  enormously  and  then  as  suddenly 
decline.  These  enormous  ducts  make  these  two  species  quite  transparent 
in  life. 

The  cestode  is  contractile  when  alive ;  it  is  from  10  to  20  cm.  long 
and  is  composed  of  from  345  to  455  proglottids,  the  average  of  14 
specimens  being  393.  In  well  expanded  specimens  the  neck  is  1.5  mm. 
long,  in  one  case  even  3  mm.  while  in  some  contracted  specimens  it  is 
only  0.3  mm.  The  diameter  varies  from  0.2  to  0.6  mm.  The  strobila 
increases  in  width  gradually  to  reach  the  maximum  width  of  1.5  mm. 
about  one-third  of  its  length  from  the  anterior  end.  From  this  point 
back  the  width  remains  about  constant.  Sexually  mature  proglottids, 
which  begin  about  the  145th  in  the  strobila,  measure  in  a  typical  speci- 
men 0.3  mm.  long  by  0.9  mm.  broad. 

The  very  large  scolex  is  nearly  globular  in  form,  being  600  to  800/x 
wide  and  700  to  950/u,  long;  eleven  specimens  which  were  measured 
averaged  709ju.  wide  by  811/x,  long.  One  finds  but  slight  indications  of 
grooves  between  the  suckers;  the  very  shallow  depressions  fade  out 
before  reaching  the  apex.  The  suckers  are  very  large,  thin-walled 
structures,  usually  collapsed  and  opening  directly  forwards.  Their 
diameter  is  300/u,  and  their  depth,  415/u.  Their  muscular  wall  is  50/n 
thick  and  the  orifice  is  50/t  across. 


12  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [362 

The  genital  pore  is  on  the  right  margin,  two-thirds  the  length  of 
the  proglottid  from  the  anterior  end.  The  cirrus  pouch  is  a  stout, 
pearshaped  structure,  155/*  long,  75/*  in  diameter  at  its  inner  end,  and 
35/*  in  diameter  at  its  outer.  The  tip  of  the  pouch  lies  dorsad  of  the 
ventral  excretory  canal  just  within  its  lateral  boundary.  Part  of  the 
vesicula  seminalis  lies  within  the  pouch;  a  much  larger  portion  lies 
without,  dorsad  of  the  ventral  excretory  duct  and  extending  mediad  of 
it.  There  is  no  prostate  gland.  The  testes  are  dorsal,  extending  nearly 
if  not  entirely  across  the  median  field,  and  into  the  lateral  field  on  the 
side  away  from  the  genital  pore.  On  the  pore  side  of  the  field  they  are 
confined  to  the  region  anterior  to  the  ovary;  on  the  opposite  side  they 
are  slightly  more  posterior  than  anterior.  They  number  43  to  57  in 
eight  proglottids  counted  and  average  about  50/*  long,  being  slightly 
smaller  in  other  dimensions.  They  break  down  early  but  each  remains 
distinct  within  its  membrane. 

The  vagina  lies  entirely  posterior  to  the  cirrus  pouch,  and  in  the 
same  dprsoventral  plane.  Its  walls  are  beset  with  glands.  Dorsad  of 
the  ventral  excretory  duct  and  extending  beyond  it  on  either  side  is  the 
very  large  receptaculum  seminis  which  is  somewhat  constricted  in  the 
middle,  giving  it  a  two-lobed  appearance.  The  ovary  consists  of  about 
15  distinct  lobes,  which  radiate  in  all  directions  from  the  point  of  origin 
of  the  oviduct.  The  posterior  lobes,  which  are  much  shorter  than  the 
rest,  extend  distad  on  the  ventral  side  of  the  transverse  commissure  to 
near  its  posterior  limit.  The  oviduct  begins  just  anterior  to  the  trans- 
verse commissure,  four-tenths  of  the  diameter  of  the  proglottid  from 
the  pore  margin. 

The  uterus  is  reticular  tho  there  is  a  tendency  for  the  tubes  not 
to  develop.  The  uterus  thus  tends  to  become  what  might  be  termed 
diffuse.  Certain  main  duqts  develop  regularly;  the  rest  of  the  cavity 
is  usally  formed  by  the  mere  expansion  of  these.  In  front  of  the  ovary, 
near  the  median  line,  the  uterus  is  represented  by  a  single  tube ;  to  the 
right  and  left  it  widens  distally  until  near  the  excretory  ducts ;  it  then 
crosses  beyond  these  ventrally.  The  later  development  of  the  uterus  is 
by  regular  outpocketing.  The  embryo  is  about  20/*  in  diameter.  It 
bears  a  large  pyriform  body  whose  length  plus  that  of  the  embryo  is 
30/*.  The  outer  embryonic  membranes  are  mostly  elongate;  when 
spherical,  the  outer  membrane  has  a  diameter  of  30  to  32/*.  The  middle 
membrane  is  loose  fitting  and  irregular.  The  uterus  of  this  species  is 
almost  identical  with  that  of  the  genus  Andrya  and  therefore  different 
from  that  of  other  closely  allied  genera.  Likewise  the  distribution  of 
the  testes  is  fundamentally  the  same.  This  species  however  disagrees 
with  other  known  Andryae  in  that  it  has  no  prostate  gland,  and  in  that 


363]  ANOPLOCEPHALID&—DOUTHITT  13 

it  has  a  well  developed  external  vesicula  seminalis.  The  extraordinary 
development  of  the  ventral  excretory  system,  moreover,  is  a  character 
of  which  one  finds  no  suggestion  in  its  nearest  allies.  These  points  argue 
strongly  for  generic  distinctness  for  this  and  the  next  species.  Whether 
to  create  for  them  a  distinct  genus  or  include  them  in  the  genus  Andrya 
is  a  question.  However  the  uterine  structure  and  testicular  distribution, 
combined  with  certain  other  undefinable  resemblances  seem  evidences 
of  sufficiently  close  generic  kinship  to  the  known  species  of  Andrya  to 
permit  the  inclusion  of  these  forms  in  this  genus.  Yet  the  diagnosis  of 
the  genus  Andrya  must  be  altered  so  as  to  admit  these  species. 

Andrya  translucida  sp.  nov. 
[Figures  14-16] 

Three  specimens  of  this  cestode  were  taken  from  a  pocket  gopher 
(Geomys  hursarius)  from  low,  semi-swampy  river  bottom  land  at 
Brainerd,  Minnesota.  The  same  gopher  yielded  also  five  specimens  of 
A.  macrocephala  to  which  it  is  evidently  closely  related.  Since  a  great 
many  gophers  were  examined  in  this  and  other  localities  (see  table, 
page  62)  it  would  seem  that  the  species  is  a  rare  parasite  of  the  gopher. 

The  total  length  of  the  specimens  at  hand  is  9  to  12  cm.  The  num- 
ber of  proglottids  is  278  to  289.  The  greatest  diameter  of  the  strobila, 
0.75  mm.,  is  reached  3  cm.  from  the  anterior  end.  Back  of  this  the 
proglottids  become  longer  and  narrower,  the  most  posterior  becoming 
attenuated  and  some  of  those  at  hand  measuring  0.3  mm.  wide  by  1.3 
mm.  long.  The  worm  is  quite  transparent  in  life,  due  to  the  enormous 
development  of  the  excretory  ducts. 

The  scolex  is  730  to  830/*  long  by  590  to  640/*  wide.  The  suckers 
stand  out  rather  prominently  and  are  separated  from  each  other  by 
distinct  grooves.  They  open  obliquely  forward,  through  orifices  about 
110/*  across.  A  definite  neck  is  present,  500  to  730/*  in  length  and  350 
to  410/*  in  diameter. 

The  genital  pore  is  usually  on  the  right  margin  but  frequently  a 
number  of  adjacent  proglottids  have  their  pores  on  the  opposite  side. 
The  pore  is  located  seven-tenths  of  the  length  of  the  lateral  margin 
from  the  anterior  end  of  the  proglottid.  The  cirrus  pouch  at  the  time 
the  sperms  begin  to  enter  is  usually  short  and  thick,  being  75/*  long  by 
40/*  broad,  and  lying  wholly  laterad  of  the  ventral  excretory  duct.  In 
succeeding  proglottids  it  lengthens  rapidly,  becoming  160/*  long  by  40/* 
wide  at  its  inner  end  which  lies  mediad  of  the  ventral  excretory  duct. 
The  vesicula  seminalis  occurs  both  within  and  without  the  pouch,  being 
represented  without  by  a  coiled  tube  which  is  about  five  times  the 


14  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [364 

diameter  of  the  vas  deferens  and  extends  forward  and  mediad  in  front 
of  the  receptaculum.  The  testes  are  usually  confined  entirely  to  the 
side  opposite  the  pore  but  two  or  three  may  lie  on  the  pore  side  of  the 
median  line;  however  they  do  not  reach  as  far  as  the  median  line  of 
the  vitelline  gland.  Likewise,  the  testes  are  nearly  all  in  the  distal 
half  of  the  proglottid.  In  both  these  features  is  seen  a  superficial 
resemblance  to  Anoplocephala.  The  testes  extend  well  across  the  ventral 
excretory  duct  laterally.  When  not  disturbed  by  the  ventral  excretory 
duct  they  extend  entirely  thru  the  medullary  portion. 

The  vagina  lies  ventral  of  the  cirrus  pouch.  Laterad  of  the  excre- 
tory ducts  its  walls  are  glandular.  Very  early,  long  before  sexual 
maturity,  it  expands  at  its  inner  end  forming  a  receptaculum  seminis. 
This  cavity  grows  and  extends  laterad  until  it  reaches  the  outer  edge 
of  the  dorsal  excretory  duct.  It  is  largest  at  its  inner  end  and  its 
diameter  decreases  gradually  laterad  so  that  it  merges  imperceptibly 
into  the  unenlarged  portion  of  the  vagina.    The  ovary  is  formed  of  about 

15  lobes  which  radiate  anteriorly,  distally,  and  laterally;  the  female 
glands  are  situated  slightly  to  the  pore  side  of  the  median  line.  The 
vitelline  gland  is  of  rather  unusual  shape,  owing  to  the  size  of  the 
median  lobe  and  its  encroachment  on  the  space  within  the  circle  of  the 
gland. 

The  uterus  is  a  very  simple  reticulum  with  its  different  branches 
wide  apart  and  clearly  distinct  from  each  other.  It  lies  anterior  to  the 
ovary  and  extends  laterad  to  beyond  the  excretory  ducts  on  either  side. 
In  front  of  the  ovary  two  or  three  transverse  tubes  are  to  be  found ;  in 
the  lateral  regions,  six  or  eight.  The  lateral  development  of  the  pro- 
glottids  is  very  peculiar,  and  apparently  abnormal,  in  that  the  ova  do 
not  pass  into  the  uterus.  In  one  specimen,  having  in  all  278  proglot- 
tids,  the  uterus  had  already  begun  to  develop  in  the  earliest  proglottid 
sectioned,  number  148.  In  proglottid  number  186  the  receptaculum 
had  reached  its  full  development  and  was  filled  with  sperms.  In  suc- 
ceeding proglottids  it  was  fully  developed  but  nearly  always  empty. 
Proglottid  number  187  showed  all  evidences  of  sexual  maturity.  The 
vesicula  and  receptaculum  are  developed,  and  the  branches  of  the 
netlike  uterus  have  become  open  tubes.  In  the  following  91  proglottids 
however  only  11  uteri  contain  eggs,  these  being  between  the  222d  and 
240th  proglottids.  The  uterus  is  distinguishable  in  all  except  the  most 
posterior,  attenuated  proglottids ;  it  never  develops  beyond  the  reticular 
stage.  In  proglottid  number  260  the  degeneration  of  the  ovary  is  nearly 
complete.  The  other  two  cestodes  at  hand  of  this  species  were  cleared 
and  examined  in  toto.     They  showed  the  same  condition ;  in  only  an 


365]  ANOPLOCEPHALIDJE—DOUTHITT  15 

occasional  uterus  are  eggs  present.  These  two  individuals  have  each 
289  proglottids;  in  all  three  cases  some  proglottids  have  been  shed. 

I  can  offer  no  certain  explanation  of  this  condition.  The  genital 
ducts  appear  normal  and  the  ova  certainly  do  not  pass  into  any  other 
organ  than  the  uterus.  There  can  hardly  be  any  doubt  therefore  that 
these  cestodes  were  incapable  of  perpetuating  themselves.  To  find  the 
end  proglottid,  or  even  several  of  the  terminal  proglottids  sterile  would 
not  be  surprising;  but  to  find  nearly  a  hundred  such  in  individuals 
that  have  already  shed  some  of  their  proglottids  is  certainly  not  to  be 
expected.  Such  a  condition  could  of  course  arise  as  a  mutant,  incapable 
of  perpetuating  itself;  in  this  case  it  could  hardly  be  thought  to  have 
arisen  from  Andrya  macrocephala,  the  nearest  known  relative ;  for  while 
these  and  that  species  resemble  each  other  in  a  general  way,  they  differ 
in  nearly  every  organ.  A  more  probable  explanation  seems  to  be  that 
the  gopher  is  not  the  normal  host  and  that  sterility  has  resulted  from 
unnatural  conditions  of  environment.  Sterility  as  a  result  of  unnatural 
environment  is  a  very  common  phenomenon  in  both  animals  and  plants ; 
and  there  is  no  reason  why  the  condition  might  not  be  found  in  ces- 
todes. Each  of  these  explanations  is  favored  by  the  fact  that  the  three 
individuals  were  all  found  in  the  same  host  and  were  of  the  same  size 
and  appearance,  making  it  appear  probable  that  they  arose  from  a 
single  infection. 

The  excretory  ducts  show  striking  features  also.  As  in  A.  macro- 
cephala the  ventral  ducts  are  enormously  developed.  In  this  case  the 
development  is  much  greater  however,  especially  that  of  the  transverse 
commissure,  so  that  often  the  proglottid  is  nearly  separated  by  the 
excretory  ducts  into  dorsal  and  ventral  parts,  the  testes  being  dorsal 
and  the  ovary  and  uterus  ventral.  The  transverse  commissures  are 
frequently  three  or  four  times  as  wide  as  the  space  between  them ;  and 
when  the  ventral  longitudinal  ducts  are  likewise  thus  developed,  the 
medullary  space  within  these  commissures  becomes  a  mere  patch  in  the 
anterior  part,  not  much  larger  than  is  necessary  to  contain  the  vitelline 
glands  and  the  female  ducts.  This  enormous  development  is  not  con- 
stant however ;  such  a  development  as  described  may  be  in  a  proglottid 
adjacent  to  one  with  ventral  ducts  60//,  in  diameter.  This  is  still  an 
unusual  size  but  is  not  greater  than  that  found  regularly  in  A.  macro- 
cephala. The  dorsal  duct  has  a  rather  sinuous  course  and  is  of  ordinary 
size.    It  lies  laterad  of  the  ventral. 

Altho  differing  in  practically  every  organ  from  A.  macrocephala, 
there  is  a  fundamental  resemblance  between  these  two  which  argues  for 
generic  relationship.  The  similarity  to  Anoplocephala  in  the  distribu- 
tion of  testes  is  interesting  but  is  not  to  be  taken  as  of  more  than  specific 


16  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [366 

import,  since  such  a  condition  is  easily  derived  from  Andrya  by  the 
suppression  of  the  most  anterior  testes. 

ANATOMICAL  CHARACTERISTICS  OF  THE  GENUS  ANDRYA 

The  additions  here  made  to  the  knowledge  of  the  genus  Andrya 
necessitate  a  radical  revision  of  our  conception  of  the  group.  These 
changes,  briefly,  are  as  follows: 

1.  The  definition  must  be  changed  so  as  to  admit  forms  in  which 
the  testes  cross  beyond  the  excretory  ducts  on  the  side  away  from  the 
genital  pore.  Also,  the  fact  that  A.  translucida  has  the  testes  only  on 
the  side  away  from  the  pore  and  mainly  in  the  distal  part  of  the  pro- 
glottid, must  be  recognized.  In  the  way  of  more  exact  definition  it 
should  be  stated  that  the  testes  are  always  chiefly  found  on  the  side 
away  from  the  pore  and  usually  mainly  anterior. 

2.  The  strictly  dextral  arrangement  of  the  genital  pores  which  is 
found  in  some  species,  must  be  given  recognition. 

3.  It  must  be  recognized  that  the  ovary  may  be  directly  median 
in  position. 

4.  It  must  be  recognized  that  the  later  development  of  the  uterus 
is  by  outpocketing,  in  a  manner  not  different  from  that  occurring  in 
Anoplocephala  and  Bertiella. 

5.  The  definition  must  be  changed  so  as  to  admit  species  without 
a  prostate  gland. 

6.  The  position  of  the  vagina  and  vaginal  pore  has  been  shown 
to  be  a  constant  character  and  should  be  given  recognition. 

7.  The  point  of  origin  of  the  oviduct  should  be  mentioned  as  a 
distinction  from  Schizotaenia. 

The  diagnosis  of  Andrya  then  becomes  as  follows : 
Anoplocephalinae,  witli  segments  much  broader  than  long,  except 
in  the  most  distal  parts  of  the  strobila.  A  single  set  of  reproductive 
organs  to  each  segment.  Genital  pores  mostly,  or  entirely  dextral. 
Genital  canals  pass  dorsal  of  longitudinal  excretory  vessels  and  nerves. 
Testes  chiefly  on  the  side  away  from  the  genital  pore,  usually  mainly 
anterior,  and  usually  extending  nearly  across  the  median  field  ante- 
riorly ;  on  the  side  away  from  the  pore  they  usually  extend  beyond  the 
excretory  ducts.  Vagina  and  vaginal  pore  strictly  posterior  to  the 
cirrus  pouch.  Female  glands  on  the  pore  side  of  the  median  line,  or 
with  the  ovary  median.  Oviduct  connects  with  ovary  beneath  the 
median  lobe  of  the  vitelline  gland.    Uterus  reticular,  becoming  saccular, 


367]  ANOPLOCEPHALIDJE—DOUTHITT  17 

then  developing  by  regular  anterior  arid  posterior  outpocketing.    Eggs 
with  pyriform  apparatus.    Adults  in  mammals. 

Type-species:    Andrya  rhopalocephala  Biehm  1881. 

Representing  the  genus  Andrya,  six  species  are  known;  besides 
those  here  studied,  these  are  rhopalocephala  and  cuniculi  (for  both  of 
which  see  Stiles,  1896).  In  addition,  Parona  (1900)  has  designated  a 
cestode  from  Dypus  aegyptius  as  A.  dipi,  with  no  word  of  description. 
As  it  is  a  pure  nomen  nudum,  it  can  not  be  taken  into  consideration. 
The  six  known  species  fall  into  two  well-defined  groups,  the  rank  of 
which  it  is  difficult  to  decide.    They  are  here  treated  as  subgenera : 

1.  Rhopalocephala-groxip.  Testes  mainly  anterior.  With  a  pe- 
dunculated prostate  gland  opening  into  the  vas  deferens  near  the  ven- 
tral excretory  vessel.  Vas  deferens  not  enlarged  outside  the  cirrus 
pouch  to  form  a  vesicula  seminalis.  Excretory  ducts  of  normal  dimen- 
sions.   Scolex  normal  in  size.    Four  species. 

A.  rhopalocephala  Riehm        A.  primordialis  Douthitt 
A.  cuniculi  R.  Blanchard        A.  communis  Douthitt 

2.  Macroce  phala-gr  o\np.  Testes  about  uniformly  anterior  and 
posterior,  or  mostly  posterior.  No  prostate  gland.  Vas  deferens  en- 
larged to  form  a  vesicula  seminalis  outside  the  cirrus  pouch.  Excretory 
ducts  enormously  developed.  Scolex  of  more  than  ordinary  size.  Two 
species : 

A.  macrocephala  Douthitt       A.  translucida  Douthitt 

Anoplocephala  wimerosa  Moniez  1880 
[Figure  17] 

Of  this  species  but  two  specimens  are  at  hand.  Admittedly,  this 
is  not  sufficient  material  to  form  a  basis  for  a  satisfactory  account  of 
anatomy ;  but  there  is  need  to  know  the  structure  of  the  older  members 
of  this  genus,  for  comparison  with  the  better  known  new  ones,  which 
do  not  fit  old  conceptions.  A.  wimerosa  has  been  known  for  33  years; 
but  it  has  received  scanty  attention,  and  several  of  the  statements  con- 
cerning it  I  find  to  be  in  error. 

The  specimens  studied  were  taken  from  Lepus  variabilis  at  Brian- 
Qon,  France,  by  R.  Blanchard  in  1891.  He  gave  several  specimens  to 
Stiles,  who,  after  he  had  studied  them,  divided  the  lot,  presenting  two 
to  the  Ward  Collection.  These  were  placed  at  my  disposal  for  study. 
One  is  a  toto  mount  and  the  other  was  sectioned.  The  account  here  given 
is  original,  except  for  the  statements  applying  to  external  form  which 
are  taken  from  the  account  of  Stiles  (1896). 


18  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [368 

Anoplocephala  wimerosa  has  a  length  of  10  mm.  and  a  breadth  of 
1.5  to  2.25  mm.,  and  contains  10  to  28  proglottids.  The  proglottids  are 
much  broader  than  long,  except  the  distal  segments,  which  rarely  be- 
come as  long  as  broad.  The  head  is  0.7  to  0.88  mm.  in  diameter,  is 
cuboid  in  form,  and  is  quite  distinct  from  the  strobila.  The  suckers 
are  0.4  mm.  in  diameter,  are  prominent,  and  open  diagonally  forward. 

As  would  be  expected  in  so  short  a  strobila,  each  proglottid  shows 
a  distinct  advance  in  development  over  the  one  before.  The  genital 
organs  are  fairly  well  formed  in  the  first  proglottid.  The  receptaculum 
seminis  begins  to  fill  with  spermatozoa  in  the  fifth.  By  the  fifteenth 
the  testes  have  practically  disappeared,  and  the  receptaculum,  vesicula, 
and  cirrus  pouch  are  enormously  distended  with  spermatozoa.  A  few 
remnants  of  the  testes  persist  until  the  eighteenth.  The  ovary  develops 
to  its  full  size  in  the  eighth  or  ninth  proglottid  and  begins  to  discharge 
eggs  into  the  uterus.  In  the  next  proglottid  this  process  is  completed 
and  the  ovary  disappears. 

The  testes,  which  are  frequently  as  much  as  90/i  long,  are  irregular 
in  shape.  They  occur  on  the  side  away  from  the  pore  and  reach  laterad 
to  beyond  the  excretory  ducts.  The  number,  in  six  proglottids  counted, 
ranged  from  31  to  42.  Not  more  than  two  testes  can  lie  in  a  row  in 
the  axis  of  the  strobila.  The  cirrus  pouch  is  enormous.  In  a  mature 
proglottid  it  is  520/1  long  and  at  its  inner  end  extends  thru  the  entire 
length  of  the  proglottid.  Its  median  end  is  occupied  by  a  portion  of  the 
vesicula  seminalis,  190/x  long.  The  portion  of  the  vesicula  outside  the 
pouch  has  a  length  in  mature  proglottids  of  100/x.  and  is  two-thirds  as 
wide.  In  older  proglottids  the  entire  vesicula  becomes  much  larger. 
The  cirrus  pouch  lies  of  necessity  almost  entirely  mediad  of  the  excre- 
tory ducts. 

The  genital  pores  are  dextral.  When  undisturbed,  the  cloaca  is 
200/i  deep.  When  the  proglottid  enters  the  period  of  sexual  activity, 
it  turns  inside  out.  In  the  third  or  fourth  segment  succeeding,  it  is 
again  drawn  back.  In  this  feature  and  in  the  enormous  size  of  the 
cloaca  and  the  cirrus  pouch  this  cestode  recalls  Schizotaenia,  especially 
S.  anoplocephaloides,  described  in  this  paper.  The  unexpanded  portion 
of  the  vagina  is  about  300/i.  long.  Throughout  its  entire  length  it  lies 
ventrad  of  the  cirrus  pouch,  as  does  also  the  vaginal  pore.  No  indica- 
tion of  glandular  structure  in  its  walls  was  observed.  The  receptacu- 
lum seminis  is  300/i.  long  in  mature  proglottids  and  at  its  median  end 
extends  from  the  anterior  to  the  posterior  border  of  the  proglottid. 
Latterly  it  narrows  regularly  and  evenly.  In  succeeding  segments  it 
becomes  enormously  distended. 


369]  ANOPLOCEPHALIDJE—DOUTHITT  19 

The  toto  mount  shows  the  whole  of  the  female  glands  to  be  situated 
slightly  to  the  pore  side  of  the  median  line;  the  other  specimen  shows 
decidedly  the  opposite  conditions  in  most  proglottids.  The  shell  gland 
is  very  indistinct  in  the  material  at  hand.  Nothing  could  be  made  out 
concerning  its  structure.  The  vitelline  gland  is  135/x  long  in  transverse 
measurement.  The  posterior  border  is  straight.  Near  the  left  end  the 
gland  measures  76/t  across  anteroposterior^ ;  it  narrows  somewhat  to- 
wards the  right  until  40/i  from  the  end ;  it  then  narrows  abruptly.  The 
ovary  in  immature  proglottids  consists  of  ten  lobes  which  radiate  ante- 
riorly and  laterally  from  a  transverse  portion.  In  its  fully  developed 
stage,  however,  no  such  divisions  can  be  recognized.  The  ovary  has  a 
width  of  420/i  and  extends  practically  through  the  proglottid  antero- 
posteriorly.  It  reaches  its  full  development  in  the  eighth  or  ninth 
proglottid  and  disappears  completely  in  the  succeeding  one. 

The  uterus  begins  as  a  simple  transverse  tube  which  seems  to  be 
confined  wholly  to  the  region  between  the  excretory  ducts.  It  develops 
mostly  by  expansion  of  the  primary  lumen,  this  occurring  first  at  the 
end  away  from  the  genital  pore.  Short  anterior  and  posterior  outpock- 
etings  are  present.  In  toto  mount  it  appears  as  if  these  pockets  are 
separated  by  a  considerable  space  from  each  other;  this  is  due  to  the 
fact  that  the  eggs  are  shrunken,  the  outlines  of  the  egg-masses  being 
taken  for  the  outlines  of  the  pockets.  As  the  uterus  becomes  filled  with 
eggs  it  appears  to  force  itself  into  the  region  ventrad  of  the  excretory 
ducts.  The  embryos  in  the  uterus  have  three  shells.  The  outer  is 
usually  somewhat  compressed ;  when  spherical  it  measures  48/u,  to  55/* 
in  diameter.  The  embryo  itself  measures  13/*  to  15.5/x,  in  diameter.  The 
length  of  the  embryo  plus  the  pyriform  body  is  17.5/t  to  21/x.  The 
appearance  of  the  horns  of  the  pyriform  body  could  not  be  made  out. 
These  measurements  would  probably  not  hold  true  for  fresher  material. 
It  was  not  possible  to  make  out  much  concerning  the  excretory  ducts. 
The  dorsal  duct  lies  laterad  of  the  ventral,  and  usually,  at  least,  in  the 
same  plane.  The  two  ducts  appear  to  be  approximately  equal  in  size. 
Transverse  commissures  are  present,  but  can  be  made  out  only  with 
difficulty. 

It  will  be  seen  that  this  species  is  without  doubt  an  Anoplocephala. 
Yet  it  disagrees  with  the  accepted  conception  of  the  genus  in  that  the 
testes  are  not  confined  to  the  median  field,  and  in  that  the  female 
glands  may  not  be  placed  to  the  pore  side  of  the  median  line.  Both  of 
these  conditions  were  found  in  Andrya,  where  they  were  supposed  not  to 
occur ;  and  the  first  was  found  also  in  another  species  of  Anoplocephala ; 
they  cannot  therefore  be  regarded  as  generic  differences.    The  diagnosis 


20  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [370 

of  the  genus  Anoplocephala  must  of  course  be  altered  to  conform;  this 
matter  is  taken  up  later  on. 

Anoplocephala  variabilis  sp.  nov. 

[Figures  18-24] 

This  cestode  has  been  found  in  Geomys  bursarius,  from  Springfield, 
Illinois,  to  Emerson,  Manitoba ;  as  yet,  no  examinations  have  been  made 
south  of  this  region.  It  seems  to  be  wholly  absent  from  the  sandhill 
regions;  at  Bemidji,  Minnesota  a  few  were  found  in  hosts  from  very 
wet,  coarse  sand  mixed  with  humus,  only  a  foot  or  so  from  the  edge  of 
a  swamp.  In  fertile  soil,  both  uplands  and  bottoms,  they  vary  in  fre- 
quency, being  in  some  localities  extremely  abundant,  and  in  others  rare 
or  apparently  absent.  Not  infrequently  as  many  as  fifty  will  be  taken 
from  a  single  host;  in  one  case  136  specimens  of  this  and  a  species  of 
Hymenolepis  were  taken  from  one  gopher.  For  distribution  and  fre- 
quency, see  the  table  on  page  62.  From  this  table  it  would  seem  proba- 
ble that  the  range  of  this  species  is  the  same  as  that  of  its  host  tho  it  is 
absent  in  many  localities  where  its  host  occurs.  The  following  descrip- 
tion is  based  on  well  expanded  specimens  from  central  Illinois. 

The  total  length  varies  from  45  to  75  mm ;  in  life  they  can  contract 
to  20  mm.  or  possibly  less.  The  proglottids  number  from  175  to  225, 
the  average  of  19  specimens  counted  being  196.  A  neck  is  usually  pres- 
ent which  is  four-fifths  the  diameter  of  the  head,  and  one-half  to  one 
and  one-half  times  as  long,  depending  upon  the  stage  of  contraction. 
The  strobila  increases  gradually  in  width  to  the  posterior  end,  being 
there  2  to  3  mm.  wide;  occasionally  the  last  few  proglottids  become 
narrower  and  longer.  The  first  proglottids  are  very  short.  Distad 
they  lengthen  rapidly,  most  of  the  proglottids  being  two-thirds  as  long 
as  broad.  In  several  specimens  especially  in  immature  regions  they 
are  longer  than  broad.  This  is  true  only  in  case  of  extreme  expansion, 
however.  The  scolex  is  irregularly  oval  in  outline  with  sometimes  a 
suggestion  of  quadrilaterality.  Ordinarily  it  is  about  275/u  long  by  300 
to  400/t  broad.  It  is  distinctly  set  off  from  the  neck  by  a  deep  circular 
groove  which  however  is  not  always  apparent.  The  suckers  stand  out 
rather  prominently,  due  to  the  presence  of  longitudinal  grooves  between 
them,  the  dorsal  and  ventral  grooves  being  deepest  and  extending  far- 
thest distad.  Opposite  the  anterior  end  of  the  suckers  the  dorsal  and 
ventral  grooves  arch  over,  the  arches  being  as  prominent  as  or  more 
prominent  than  the  suckers.  These  arches  are  the  ends  of  a  transverse 
bar  which  stands  out  prominently  as  the  most  anterior  part  of  the 


371]  ANOPLOCEPHALIDJE—DOUTHITT  21 

scolex.  In  frontal  section  this  bar  shows  as  a  projecting  tip  in  the 
median  line.  In  some  specimens  however  the  bar  and  arches  are  not 
recognizable.  The  grooves  never  extend  to  the  tip.  The  suckers  open 
obliquely  forward,  at  an  angle  of  about  45  degrees  from  the  axis  of  the 
strobila.  The  cuticula  surrounding  the  orifice  may  be  a  simple  fold  or 
there  may  be  as  many  as  four  folds  of  varying  prominence. 

The  genital  anlagen  are  represented  by  deeply  staining  cells  even 
before  strobilization  is  evident.  The  organs  begin  to  take  on  recogniza- 
ble form  about  the  45th  proglottid,  and  eggs  begin  to  pass  into  the 
uterus  about  the  75th.  The  genital  pore  is  on  the  right  margin,  two- 
thirds  of  the  length  of  the  proglottid  from  the  anterior  end  and  con- 
siderably nearer  the  dorsal  than  the  ventral  surface.  In  some  speci- 
mens it  is  everted  during  sexual  activity.  The  cirrus  pouch  is  200/x 
long  by  75/x  wide,  and  pear-shaped.  It  extends  well  across  the  excretory 
ducts,  dorsad  of  them.  The  cirrus  is  not  spiny.  Part  of  the  vesicula 
seminalis  occupies  the  median  end ;  outside  the  pouch  the  vesicula  bends 
upon  itself  twice,  enlarging  greatly  at  its  inner  end.  From  here  the 
vas  deferens  takes  a  somewhat  convoluted  course  across  the  proglottid. 
The  testes  shrink  and  apparently  break  down  very  early  so  that  indi- 
vidual recognition  is  difficult.  The  number  varies  from  60  to  85;  they 
are  about  65fi  long  antero-posteriorly  and  usually  somewhat  smaller  in 
other  dimensions.  They  occur  in  the  side  away  from  the  pore,  extend- 
ing from  the  median  line  to  the  nerve  trunk  and  from  ten  to  twenty  lie 
outside  the  excretory  ducts.  They  extend  forward  to  slightly  beyond 
the  uterus  and  in  the  central  part  of  their  field  entirely  through  the 
medullary  portion.  In  expanded  specimens  it  is  clear  beyond  doubt 
that  they  are  to  be  considered  distal  in  occurrence,  rather  than  proximal. 
The  vaginal  pore  is  on  the  ventral  surface  of  the  cloaca.  The  vagina 
lies  ventral  of  the  cirrus  pouch,  and  at  its  median  end  usually  some- 
what posterior  to  it.  Its  walls  are  glandular.  Just  within  the  excretory 
canals  is  the  receptaculum  seminalis,  which  is  at  first  clearly  divisible 
into  a  small  lateral  and  a  larger  median  portion,  both  of  which  are 
about  globular  in  form.  Within  the  circle  of  the  vitelline  gland  the 
receptaculum  becomes  a  short,  thickwalled  tube,  which  joins  the  oviduct. 

The  ovary  lies  mostly  to  the  pore  side  of  the  median  line.  It 
consists  of  a  bar  in  front  of  the  vitelline  gland,  turning  distad  beneath 
the  median  end  of  the  latter,  with  lobes  radiating  in  all  directions  in 
the  longitudinal  plane  except  directly  backwards.  The  oviduct  arises 
from  beneath  the  median  end  of  the  vitelline  gland  mediad  of  the  cen- 
tral axis  of  the  ovary.  It  takes  a  fairly  definite,  sinuous  course  laterad, 
then  proximad,  to  join  the  vagina.  The  common  duct  takes  an  irregular 
course  to  where  it  passes  into  the  shell  gland.    The  vitelline  gland  is  a 


22  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [372 

large,  horseshoe-shaped  body,  just  to  the  pore  side  of  the  median  line. 
Its  duet  arises  from  near  the  median  line  of  the  gland.  It  can  be 
traced  inside  the  gland  as  an  open  passage  which  divides  and  goes  to 
the  two  lobes.  When  it  has  emerged  from  the  shell  gland  the  uterine 
duct  passes  forward  and  median  with  several  windings  which  are  fairly 
constant  in  different  individuals,  to  join  the  uterus  near  the  median 
line.  The  uterus  is  at  first  a  narrow,  sinuous  transverse  tube,  just 
below  the  upper  limit  of  the  ovary  and  testes.  At  either  end  it  crosses 
beyond  the  excretory  ducts  ventrally,  and  turns  distad  beyond  them. 
Just  before  the  eggs  begin  to  enter  the  uterus  expands  considerably. 
The  later  development  is  by  outpocketing,  the  walls  of  the  adjacent 
pockets  touching.  The  anterior  pockets  number  28  to  40  being  usually 
about  35;  the  posterior  pockets,  22  to  33,  being  usually  about  30. 

The  embryo  has  three  membranes.  The  outer  is  spherical,  30  to 
35/i  in  diameter;  the  middle  is  loosefitting  and  irregular.  The  embryo 
itself  is  12  to  13/x  in  diameter.  It  bears  a  well  developed  pyriform 
apparatus.  The  dorsal  excretory  duct  lies  dorsal  of  the  ventral  in 
young  proglottids.  As  the  proglottid  ripens  it  passes  laterad  and  usu- 
ally ventrad  also.  At  sexual  maturity  the  dorsal  duct  is  usually  lateral 
but  it  may  be  both  strictly  dorsal  and  strictly  lateral  in  different  parts 
of  the  same  proglottid.    The  transverse  commissure  is  sinuous. 

The  foregoing  description,  as  already  stated,  is  based  on  well 
expanded  material  from  central  Illinois.  In  the  material  from  the  same 
localities,  however,  are  a  number  of  very  much  contracted  specimens 
which  are  so  different  from  the  forms  already  described  that  they 
would  no  doubt  have  been  classed  as  separate  species  if  only  the  pre- 
served specimens  were  known.  The  main  differences  from  the  type 
already  described  are  as  follows: 

The  length  is  30  to  33  mm.  The  proglottids  are  all  very  short  and 
broad.  Mature  proglottids  are  0.325  mm.  long  by  4.5  mm.  broad.  The 
suckers  open  directly  forward.  The  scolex  is  half-moon  shaped,  and  is 
not  broader  than  the  first  part  of  the  strobila.  The  genital  pore  is 
situated  midway  on  the  right  margin  nearer  the  dorsal  than  the  ventral 
surface.  The  ovary  extends  from  the  ventral  excretory  canal  on  the 
pore  side  to  not  far  from  the  corresponding  canal  on  the  opposite  side. 
The  oviduct  arises  from  the  median  line  of  the  proglottid,  half  its  length 
lying  mediad  of  the  vitelline  gland.  The  transverse  commissure  of  the 
excretory  duct  is  not  sinuous.  The  dorsal  excretory  duct  is  always 
lateral  to  the  ventral,  except  in  the  head  region  and  vicinity.  In  mak- 
ing comparisons  of  these  two  types,  the  reader  should  refer  to  figures 
18,  19,  22,  and  23. 


373]  AN0PL0CEPHAL1DAZ—D0UTHITT  23 

Perhaps  the  most  striking  of  these  differences  is  seen  in  the  position 
of  the  genital  pore.  This  is  explained  however  when  one  notes  that  in 
the  elongate  proglottid  the  portion  of  the  body  wall  anterior  to  the 
genital  pore  lies  much  nearer  the  longitudinal  muscles,  and  more  nearly 
parallel  to  them,  than  does  the  portion  posterior.  As  a  consequence  the 
anterior  part  would  be  shortened  much  more  than  the  posterior  in  the 
process  of  contraction.  That  such  is  the  case  is  shown  by  the  fact  that 
in  contracted  specimens  the  cuticula  is  much  thicker  anterior  to  the 
pore  and  is  thrown  into  many  fine  wrinkles. 

In  order  to  make  sure  that  these  were  really  of  the  same  species, 
experiments  were  performed  with  the  object  of  trying  to  produce  the 
two  types  at  will.  Complete  success  was  attained  with  the  simplest 
methods  of  treatment.  Living  specimens,  just  removed  from  the  host, 
were  cut  each  into  two  pieces;  one  was  placed  in  a  dish  of  water,  the 
temperature  of  which  was  estimated  to  be  about  30°  C  ,•  the  other  in  a 
dish  of  about  37°,  care  being  taken  that  the  portions  from  different 
worms  were  not  confused.  Those  in  the  warmer  dish  soon  expanded, 
and  while  in  this  condition  they  were  fixed  in  corrosive  sublimate.  Those 
in  the  colder  water  contracted,  however,  and  could  not  be  induced  to 
expand.  When  sectioned  the  portions  from  warm  water  were  found  to 
be  without  exception  of  the  type  first  described,  and  those  from  the 
cold  water  of  the  second  type.  This  simple  experiment  shows  the 
necessity  in  comparing  cestodes  of  taking  into  account  the  state  of 
contraction  of  the  proglottid. 

These  descriptions,  as  already  stated,  apply  to  specimens  from 
central  Illinois.  As  one  passes  northward,  however,  it  is  found  that  the 
individuals  grow  steadily  smaller,  both  in  bulk  and  number  of  proglot- 
tids.  In  anatomical  features  only  one  difference  was  discerned:  the 
testes  become  very  regularly  fewer;  apparently  their  size  remains  about 
the  same,  tho  their  irregular  shape  does  not  permit  of  exact  determina- 
tion on  this  point.  The  most  conspicuous  difference  between  the  worms 
from  different  localities  is  in  size,  those  from  the  north  being  only  about 
half  the  length  and  breadth  of  those  from  the  south.  The  other  differ- 
ences are  illustrated  in  the  following  table.  The  numbers  in  parenthe- 
ses indicate  the  numbers  of  individuals  or  proglottids  examined. 


24 


ILLINOIS    BIOLOGICAL    MONOGRAPHS 


[374 


Locality 


Latitude 


Number  of 

Proglottid 

Number  of      in  which      Number  of 

Proglottids     Eggs  first         Testes 

in  uterus 

196      (19)     75-8i   (6)        68.9  (10) 


190.5  (2) 

76 

(1) 

53-4  (5) 

154-8  (19) 

65,68(2) 

45-5  (10) 

147   (2) 

63 

(1) 

144-7  (3) 

54 

(1) 

129   (3) 

56 

(r) 

43-3  (7) 

134-5  (2) 

57 

(1) 

40.2  (10) 

Lincoln,  Springfield 

and  Clinton,  Illinois 400 

Minneapolis,  Minnesota  450 

Brainerd,  Minnesota  450  21' 

Wahpeton,  North  Dakota 450  21' 

Bemidji,  Minnesota  47°  28'  30' 

Thief  River  Falls,  Minnesota.  48°  7' 
Emerson,  Manitoba  49° 


In  judging  from  these  figures  it  must  be  borne  in  mind  that  while 
Emerson,  Manitoba  is  considerably  farther  north  than  Bemidji  and 
Brainerd,  Minnesota,  it  is,  nevertheless,  in  the  transitional  zone  while 
they  are  in  the  boreal.  It  will  be  observed  from  this  table  that  while 
there  is  a  considerable  difference  between  the  extremes,  the  intergrada- 
tions  are  sufficiently  regular  to  destroy  the  validity  of  these  differences 
as  specific  characteristics.  They  are  therefore  here  classed  as  one  spe- 
cies, with  the  specific  name  "variabilis".  The  northern  forms  are  fur- 
ther recognized  as  a  variety,  with  the  name  Anoplocephala  variabilis 
"borealis. 


Anoplocephala  infrequens  sp.  nov. 

[Figures  25  to  27] 

This  cestode  was  found  rather  sparingly  in  Geomys  bursarius  in 
northern  Minnesota  and*  just  across  the  Canada  line,  at  Emerson,  Mani- 
toba. Recently  also,  through  Professor  Ward  I  have  received  from 
Professor  Young  a  cestode  taken  from  Evotomys  sp.  at  Grand  Forks, 
North  Dakota,  which  is  identical  with  these.  In  all,  ten  specimens  were 
secured  from  the  gophers.  All  were  taken  from  the  last  inch  of  the 
small  intestine  or  the  adjacent  part  of  the  large  intestine,  were  un- 
attached, and  showed  no  signs  of  life.  Whether  or  not  they  belong  this 
far  back  cannot  be  said  positively;  but  the  fact  that  three  were  imma- 
ture and  that  all  but  one  were  in  good  condition  when  found  would 
seem  to  indicate  that  they  had  not  died  and  drifted  back.  The  follow- 
ing description  was  based  upon  three  sectioned  and  three  adult  alcoholic 
specimens.  Later,  the  specimen  from  Evotomys  was  sectioned  and 
compared;  no  significant  points  of  disagreement  with  the  account  here 
presented  were  discovered. 


375]  ANOPLOCEPHALIDAZ—DOUTHITT  25 

The  total  length  is  12.5  to  20  mm.  The  number  of  proglottids  is 
61  to  72.  There  is  no  neck.  The  first  proglottid  is  about  40/i  long,  and 
generally  about  five-sixths  the  diameter  of  the  head.  The  strobila  may 
increase  rapidly  in  width  in  the  first  few  proglottids,  or  the  fourth  or 
fifth  may  be  four-fifths  the  diameter  of  the  first.  Mature  proglottids 
measure  1.5  mm.  wide  by  145/i.  long.  The  maximum  width  of  2.5  to  5 
mm.  is  reached  about  7  mm.  from  the  posterior  end.  Beyond  this  the 
proglottids  usually  narrow  rapidly  and  lengthen  somewhat.  The  pos- 
terior free  border  of  each  proglottid  overlaps  about  half  of  the  suc- 
ceeding proglottid. 

The  scolex  is  625  to  750/u,  broad  near  its  anterior  end.  It  narrows 
distad,  passing  abruptly  into  the  first  proglottid.  It  is  nearly  square 
in  cross  section  with  only  slight  indication  of  grooves  between  the 
suckers.  On  the  anterior  end  grooves  occur  between  the  suckers,  fading 
out  just  before  the  apex  is  reached.  There  is  no  indication  of  the 
circular  groove  found  in  the  last  species.  The  very  large  suckers  open 
directly  forward.  Right  and  left  suckers  touch  in  the  median  line  of 
the  scolex,  and  reach  very  nearly  to  the  lateral  cuticula  (see  Fig.  26). 
The  mouth  is  70  to  90/x  across.    The  muscular  wall  is  55  to  75/x  thick. 

The  genital  anlagen  appear  in  the  first  proglottid.  The  receptacu- 
lum  seminis  fills  in  the  17th  or  18th,  and  eggs  are  found  in  the  uterus 
in  the  21st  or  22nd.  The  genital  pores  are  located  midway  on  the 
right  margin  and  distinctly  nearer  the  dorsal  than  the  ventral  surface. 
The  cirrus  pouch  extends  first  proximo-mediad  and  then  bends  directly 
mediad,  not  enlarging  beyond  the  bend.  Its  inner  end  lies  well  within 
the  excretory  ducts.  The  passage  of  the  cirrus  is  tortuous.  In  the 
median  end  of  the  pouch  is  a  portion  of  the  vesicula  seminalis  which 
later  enlarges  to  fill  the  entire  pouch.  Outside  the  pouch  is  a  second 
portion  of  the  vesicula.  The  testes  shrink  and  apparently  break  down 
very  early.  They  are  50  to  60  in  number  and  are  70/i  long,  being 
usually  somewhat  longer  than  broad.  Nearly  all  lie  between  the  left 
ventral  excretory  vessel  and  the  vitelline  gland,  but  not  infrequently 
a  testis  will  lie  wholly  or  in  part  laterad  of  the  excretory  vessel  and 
two  or  three  may  lie  dorsad  of  the  anterior  part  of  the  vitelline  gland. 
They  extend  entirely  thru  the  medullary  portion  dorsoventrally. 

The  vaginal  pore  is  on  the  ventral  surface  of  the  cloaca.  The 
vagina  lies  ventrad  to  the  cirrus  pouch  and  partly  behind  it.  Its  walls 
are  highly  glandular.  The  receptacular  seminis  is  large  and  elongate, 
reaching  laterad  to  the  ventral  excretory  vessel.  A  small  part  at  the 
lateral  end  is  indistinctly  constricted  off  from  the  median  portion, 
making  it  somewhat  bilobed  The  oviduct  arises  just  anterior  to  the 
lower  median   corner  of  the  vitelline   gland  and   passes  laterad  and 


26  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [376 

proximad  with  several  turns  over  the  ventral  surface  of  the  latter. 
The  vitelline  duct  extends  as  a  large  channel  into  each  lobe  of  the  gland. 
The  uterine  duct,  after  passing  thru  the  voluminous  shell  gland,  takes 
a  crooked  but  regular  course  forward  and  mediad  to  the  uterus.  The 
very  slender  transverse  uterus  extends  laterad  at  either  end  to  beyond 
the  nerve  trunks.  Development  is  by  extension  and  outpocketing  in 
the  usual  manner.  The  anterior  pockets  number  44  to  51;  the  poste- 
rior, 28  to  34.  The  ovary,  in  individuals  from  the  gopher,  is  small, 
just  reaching  to  the  median  line  on  the  left,  and  not  reaching  the 
excretory  duct  on  the  right.  In  the  specimen  from  Evotomys,  the 
uterus  is  somewhat  larger.  The  ovary  does  not  cross  the  uterus 
anteriorly. 

The  embryo  is  typical.  The  outer  membrane  is  usually  elongate, 
or  of  other  shape,  due  apparently  to  pressure.  When  spherical,  it 
measures  39  to  43/t  in  diameter.  The  middle  membrane  is  loosefitting 
and  irregular.  The  embryo  itself  measures  12jt  in  diameter.  It  has  a 
typical  pyriform  apparatus  whose  length,  plus  that  of  the  embryo,  is 
19  to  22/i. 

The  dorsal  excretory  duct  lies  laterad  of  the  ventral.  The  inner 
duct  curves  very  strongly  laterad  in  the  middle  of  the  proglottid;  the 
curve  of  the  outer  is  much  less.  The  transverse  duct  is  sinuous  in 
dorsoventral  plane. 

THE  GENUS  ANOPLOCEPHALA 

The  genus  Anoplocephala  has  served  as  a  refuge  for  inadequately 
described  species  throughout  its  history.  Most  of  the  older  species  of 
the  family  Anoplocephalidae  have  been  placed  here  at  one  time  or 
another;  as  they  have  become  better  known,  they  have  been  removed 
one  by  one  to  other  genera.  Many  of  the  names  given  have  also  been 
shown  to  be  synonyms.  Recently,  von  Janicki  (1910),  studying  the 
cestodes  from  the  hyrax,  found  that  five  of  these  supposed  species  did 
not  belong  even  to  the  subfamily.  Deiner  (1912)  has  recently  given 
an  excellent  and  thoro  description  of  "Anoplocephala"  magna  Murie; 
for  reasons  stated  on  page  40  I  have  transferred  this  species  to  the 
genus  Schizotaenia.  I  have  made  the  same  disposal  of  Anoplocephala 
(Taenia)  gigantea,  whose  anatomy  MacCallum  and  MacCallum  (1912) 
have  recently  investigated  in  a  thoro  manner. 

Unfortunately,  however,  many  of  the  species  assigned  to  the  genus 
still  remain  practically  unknown;  and  more  unfortunately,  still  other 
inadequately  described  species  are  being  added.  Thus,  Mello  (1912) 
thrusts  upon  the  world  A.  minima,  without  giving  any  information 


377]  ANOPLOCEPHALIDAZ—DOUTHITT  27 

whatever  except  to  external  form;  and  Galli-Valerio  (1905)  burdens 
science  with  A.  dentata,  with  but  two  lines  of  information  as  to  internal 
anatomy  and  these  lines  devoted  to  non-essential  details.  There  is  no 
proof  that  either  of  these  two  species  belongs  even  to  the  subfamily; 
and  indeed,  the  only  important  fact  made  known  concerning  A.  minima, 
that  the  genital  pores  are  regularly  alternate,  indicates  that  it  does  not 
belong  to  the  genus  Anoplocephala. 

If  consideration  be  given  to  these  half-starved  species,  no  anatomi- 
cal generalization  concerning  the  genus  Anoplocephala  is  possible.  Thus, 
"A."  spatula  has  the  testes  extending  across  the  median  field;  "A." 
omphalodes  has  the  genital  pores  irregularly  alternate,  the  vaginal  pore 
posterior,  and  the  testes  sometimes  extending  across  the  median  field; 
and  "A."  minima  has  the  pores  regularly  alternate.  For  about  half 
the  species  assigned  to  the  genus,  no  statement  whatever  is  made  con- 
cerning the  uterus.  There  seems  no  alternative,  then,  if  one  would 
attempt  any  generalizations,  but  to  base  them  only  on  those  species 
whose  position  in  this  genus  is  assured,  and  to  ignore  seeming  contra- 
dictions coming  from  the  inadequately  described  species  which  are 
placed  provisionally  in  the  genus. 

The  diagnosis  of  the  genus  should  be  changed  so  as  to  admit  species 
in  which  the  testes  extend  beyond  the  excretory  ducts  on  the  side  away 
from  the  genital  pore.  Also,  it  should  state  that  the  vagina  and  vaginal 
pore  are  ventral  to  the  cirrus  pouch.  It  seems  probable  that  in  all  the 
proximal  end  of  the  oviduct  lies  beneath  the  median  lobe  of  the  vitelline 
gland ;  but  information  is  needed  concerning  more  species  on  this  point. 

The  diagnosis  of  the  genus  Anoplocephala  thus  becomes  as  follows. 

Anoplocephalinae,  with  segments  generally  much  broader  than 
long,  occasionally  longer  than  broad.  A  single  set  of  reproductive  or- 
gans in  each  segment.  Genital  pores  unilateral.  Genital  canals  pass 
on  the  dorsal  side  of  the  longitudinal  excretory  vessels  and  nerve. 
External  vesicula  seminalis  present.  Testes  on  the  side  away  from  the 
genital  pore,  sometimes  extending  laterad  of  the  nerve  trunk.  Vagina 
and  vaginal  pore  ventral  to  the  cirrus  pouch.  Female  glands  to  the 
pore  side  of  the  median  field.  Uterus  a  transversely  elongated  sac,  with 
pocket-like  appendages,  anteriorly  and  posteriorly.  Eggs  with  well 
developed  pyriform  apparatus.     Adults  in  mammals. 

Type-species :    Anoplocephala  perfoliata  Goeze  1782. 


28  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [378 

The  following  species  may  be  assigned  with  certainty  to  the  genus 
Anoplocephala. 

A.  perfoliata  Goeze  A.  magna  Abildgaard 

(syn.,  A.  plicata  &  A.  zebrae) 
A.  globiceps  Diesing  A.  mamttlana  Mehlis 

A.  wimerosa  Moniez  A.  variabilis  Douthitt 

A.  infrequens  Douthitt 

The  following  species  are  not  well  enough  known  to  allow  of  generic 
determination,  but  should  be  left  here  for  lack  of  better  disposal. 

A.  inermis  von  Linstow  A.  blanchardi  Moniez  V 

(syn.,  A.  arvicolae)  A.  dentata  Galli-Valerio 

A.  omphalodes  Hermann  A.  paronai  Moniez 

A.  restrict  a  Railliet  A.  spatula  von  Linstow 

A.  transversaria  Krabbe 

Schizotaenia  americana  Stiles  1895 
[Figures  28,  29] 

Stiles  in  1895  gave  a  short  description  of  this  cestode  and  placed 
it  in  the  genus  Andrya.  The  next  year  he  transferred  it  to  the  genus 
Bertia,  (now  Bertiella).  In  1906  von  Janicki,  having  proposed  the 
genus  Schizotaenia,  included  this  form  and  Bertiella  americana  leporis 
in  that  genus,  basing  his  action  mainly  upon  the  distribution  of  testes, 
since  the  description  of  Stiles  was  noncommittal  or  erroneous  on  other 
points  that  might  be  of  importance.  The  results  of  the  present  studies 
justify  completely  von  Janicki 's  disposal. 

In  1906  Cohn  expressed  it  as  his  opinion  that  this  cestode  was  the 
one  described  by  Leidy.  (1855)  as  Taenia  laticephala,  and  that  the 
specific  name  "americana  Stiles"  should  be  dropped  as  a  synonym. 
I  cannot  agree  that  the  evidence  justifies  such  a  conclusion.  With  full 
consideration  for  the  ability  of  cestodes  to  contract  and  expand,  it  is 
a  severe  strain  on  one's  credulity  to  conceive  how  a  cestode  1.3  inches 
long  and  2.8  lines  broad  could  at  will  become  9  inches  long,  and  %  of 
a  line  broad;  and  how  a  proglottid  12  times  as  broad  as  long  could 
become  square.  Moreover,  Leidy  states  that  the  width  of  the  neck  in 
the  forms  at  his  disposal  is  14  of  a  line,  or  one-half  the  diameter  of  the 
head.  These  specimens  have  no  neck  and  strobilzation  is  very  con- 
spicuous even  in  the  first  proglottids.  Leidy  states  that  the  anterior 
proglottids  are  oblong-square.  If  this  means  that  they  were  longer  than 
broad,  we  have  a  condition  that  is  unprecedented  in  the  Anoplocephali- 
dae;  if  it  means  that  they  were  broader  than  long,  then  Leidy 's  speci- 


379]  ANOPLOCEPHALIDJE—DOUTHITT  29 

mens  must  have  been  composed  of  at  least  225  proglottids,  more 
probably  of  350  or  400  proglottids. 

The  fact  that  Leidy's  and  Stiles'  material  were  from  the  same 
host,  the  only  point  that  Cohn  seems  to  have  taken  into  consideration, 
means  nothing.  I  have  taken  16  species  of  cestodes  from  Geomys,  and 
have  not  exhausted  the  field;  Stiles  (1896)  has  reported  still  another. 
These  species  represent  seven  genera,  and  three  families.  The  balance 
of  evidence  seems  to  be  in  favor  of  Stiles'  view  that  Leidy's  species  was 
a  species  of  the  genus  Davainea. 

Cohn  criticise  Stiles  for  the  inadequateness  of  his  description;  but 
with  sectioned  material  before  him  he  failed  to  add  anything  of  impor- 
tance to  our  knowledge.  Incidentally  he  mentions  that  the  female 
glands  are  situated  at  a  considerable  distance  from  the  median  line, 
which  proves  that  he  did  not  have  before  him  Stiles'  species  but  had 
probably  Schizotaenia  variabilis,  the  next  species  discussed  in  this 
paper. 

There  were  available  for  study  specimens  from  Stiles'  material 
loaned  by  the  Bureau  of  Animal  industry;  others  were  placed  at  Pro- 
fessor Ward's  disposal  by  Professor  M.  J.  Elrod,  the  original  collector 
of  the  material;  also,  from  the  Bureau  of  Animal  Industry  a  specimen 
from  a  porcupine  taken  at  Mayfield,  Michigan.  The  first  lot  is  from 
Erethizon  epixanthus,  from  Snake  River,  Wyoming;  the  second  is  from 
Erethizon  dorsatus.  I  have  also  specimens  (No.  1502,  B.  A.  I.  Coll.) 
identified  by  Stiles  as  this  species;  examination  has  shown  that  they 
are  distinct. 

External  form  (Stiles  (1896) — "Strobila  attains  33  mm.  in  length 
by  6  mm.  in  breadth  and  contains  about  90  segments,  the  oldest  of 
which  are  8  mm.  long.  Head  unarmed,  measures  0.6  mm.  broad  by  0.38 
mm.  long  by  0.32  mm.  thick,  and  is  nearly  rectangular  in  apex  view. 
The  neck  is  absent,  and  the  head  is  frequently  retracted  in  the  body, 
as  in  Drepanidotaenia  lanceolata.  Suckers  round,  0.176  mm.  in  diame- 
ter, open  anteriorly". 

The  genital  organs  alternate,  in  the  specimens  at  hand,  with  per- 
fect regularity  from  right  to  left.  The  genital  anlagen  are  visible  in  the 
first  proglottid.  The  specimen  from  Mayfield,  Mich.,  shows  the  70th 
proglottid  still  immature;  apparently,  maturity  is  reached  about  the 
80th.  No  other  specimen  at  hand  gave  information  as  to  this  point. 
One  must  conclude  from  this  condition,  either  that  Elrod 's  material 
was  all  immature  since  it  shows  the  total  number  of  proglottids  to  be 
about  90,  or  that  the  species  is  larger  when  inhabiting  Erethizon  dorsa- 
tus. The  latter  conclusion  seems  more  probable,  since  some  of  the 
detached  proglottids  of  the  Elrod  material  are  ripe. 


30  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [380 

The  genital  cloaca  could  not  be  studied  as  carefully  as  desired,  on 
account  of  its  habit  of  turning  inside  out  at  the  approach  of  sexual 
maturity.  A  full  comparison  of  the  organ  with  that  of  Schizotaenia 
anoplocephaloides,  to  be  described  later,  is  therefore  not  possible.  They 
agree  however  in  all  points  that  could  be  made  out.  In  both  the  cloaca 
everts  at  the  approach  of  sexual  maturity ;  and  in  both  it  is  drawn  back 
at  the  close  of  sexual  activity,  but  in  none  of  the  sectioned  material  at 
hand  does  it  come  back  completely.  In  both  the  cloaca  is  very  large 
tho  in  the  present  species  it  is  not  so  extraordinarily  large  as  in  the 
other.  The  proglottids  sectioned  do  not  show  whether  the  cloaca  is 
divided  into  two  parts,  as  in  S.  anoplocephaloides.  The  pore  is  located 
about  midway  on  the  margin. 

The  cirrus  pouch  is  quite  large,  tho  small  as  compared  with  that 
of  S.  anoplocephaloides.  When  the  cloaca  is  not  everted,  about  a  third 
of  it  lies  mediad  of  the  excretory  ducts ;  when  everted  it  reaches  usually 
just  to  the  nerve  trunk.  It  is  500-625/x  long  by  185-210/u,  wide.  It  is 
somewhat  pear-shaped  in  outline,  the  inner  half  being  about  uniform  in 
diameter.  Laterad  of  the  middle  it  tapers  sharply  to  end  in  a  slender 
neck  40/a  in  diameter  and  150/x  long.  The  cirrus  is  spiny.  Extruded 
cirri  however  are  smooth,  from  which  it  would  seem  that  the  attachment 
of  the  hooks  is  very  light.  This  probably  explains  the  apparent  absence 
of  spines  observed  by  Stiles  for  S.  americana  leporis.  The  inner  end 
of  the  cirrus  pouch  is  occupied  by  a  portion  of  the  vesicula  seminalis. 
Outside  the  cirrus  pouch  the  vesicula  is  about  80/x  in  diameter  in 
mature  proglottids  and  extends  mediad  and  proximad  for  a  distance 
of  about  850/x,  coming  to  lie  here  near  the  anterior  end  of  the  proglottid. 
It  then  curves  usually  to  extend  mediad  and  distad  for  a  distance  of 
about  225/i  where  it  ends  in  the  slender  vas  deferens.  The  vasa  effer- 
entia  are  in  a  large  part  plainly  visible.  The  vesicula  seminalis  if 
closely  examined  is  seen  to  be  somewhat  sinuous;  but  the  effect  is  that 
of  a  straight  tube  taking  the  course  mentioned.  At  its  extreme  inner 
end  just  at  this  juncture  with  the  vas  deferens  it  has  glandular  walls. 
The  testes  are  spherical  and  65/t  to  80/t  in  diameter.  They  are  all 
dorsal  and  extend  from  excretory  duct  to  excretory  duct  in  the  distal 
end  of  the  proglottid.  The  number  was  found  to  be  about  70  which 
agrees  with  the  account  of  Stiles. 

The  vagina  and  vaginal  pore  are  anterior  to  the  cirrus  pouch.  For 
about  the  first  185/i  of  its  length  the  vagina  is  a  very  small,  heavy- 
walled  tube  which  is  perfectly  distinct  when  once  recognized.  Beyond 
this  point  it  is  very  indistinct  and  usually  can  be  traced  only  by  the 
vacuolated  appearance  of  its  course.  In  a  few  cases,  however,  it  can 
be  seen   very  indistinctly  anterior  to  the  cirrus  pouch,   crossing  the 


381]  AN0PL0CEPHALID2E—D0UTHITT  31 

vesicula  seminalis  just  beyond  the  pouch  and  then  proceeding  mediad 
to  the  receptaculum  seminalis.  Just  before  entering  the  receptaculum 
it  again  becomes  distinct.  Thus  in  position  only  does  it  resemble  the 
conspicuous,  glandular  vagina  of  S.  anoplocephaloides.  The  rather 
small,  nearly  globular  receptaculum  seminis  lies  just  anterior  to  and 
laterad  of  the  vitelline  gland  partly  overlapping  it. 

The  female  glands  are  but  slightly  displaced  from  the  median  line. 
In  proglottids  4  to  5  mm.  wide  the  median  axis  of  the  shell  gland  is 
55/x  to  175/x  distant  from  the  median  line.  The  female  organs  of  the 
right  and  left  sides  therefore  overlap  largely.  The  transverse  diameter 
of  the  ovary  is  about  1.3  mm.  It  is  composed  of  a  transverse  portion 
in  front  of  the  vitelline  gland  from  which  numerous  small  lobes  radiate 
in  all  directions  except  towards  the  vitelline  gland.  To  either  side  of 
the  vitelline  gland  the  lateral  portions  turn  distad  and  extend  to  near 
the  transverse  commissure  of  the  excretory  duct.  The  ova  measure 
12/a  to  15/*  in  diameter.  The  oviduct  is  short;  its  attachment  to  the 
ovary  is  located  anterior  to  the  central  axis  of  the  vitelline  gland.  This 
latter  gland  is  of  the  regular  Anoplocephaline  form,  being  composed 
of  a  small  lateral  and  a  large  median  lobe.  Here,  however,  is  a  clear 
approach  to  the  peculiar  mulberry-like  form  found  in  the  next  species 
and  in  S.  hagmanni.  The  two  lobes  approach  each  other  anteriorly, 
thus  restricting  the  space  between  them  and  making  the  gland  more 
compact  and  show  an  indication  of  the  division  of  the  gland  into  many 
small  radiating  lobes.  The  transverse  diameter  of  the  gland  is  400/a. 
The  shell  gland  is  90ii  in  diameter;  it  needs  no  further  description. 

The  uterus  in  the  earliest  stages  observed  is  a  continuous  sheet  of 
tissue  extending  through  nearly  the  whole  of  the  median  field,  except 
for  the  portion  occupied  by  the  ovary  and  vitelline  gland.  The  two 
lateral  portions  are  connected  by  a  narrow  strip  above  the  ovary.  There 
is  abundant  indication  of  thickening  of  the  uterine  tissue  into  definite 
lines,  thus  simulating  a  network  such  as  is  found  in  Moniezia.  These  do 
not  develop  into  open  tubes,  however,  so  it  is  not  really  a  reticulum. 
For  a  discussion  of  this  type  of  uterus  see  under  8.  anoplocephaloides. 
The  condition  of  the  material  unfortunately  does  not  permit  the  devel- 
opment of  this  organ  to  be  worked  out  in  full  detail.  In  its  fully  devel- 
oped stage  it  is  divided  up  into  a  great  many  small  compartments,  each 
containing  several  eggs,  similar  to  the  condition  described  later  for 
Moniezia  expansa.  It  seems,  therefore,  that  the  uterus  does  not  break 
down  into  egg-capsules  after  the  manner  of  the  Linstowinae,  but 
that  the  condition  is  due  to  unequal  development  of  the  original  cavity 
at  different  points.  In  its  fully  developed  stage  the  uterus  is  practically 
confined  to  the  median  field  but  a  small  branch  extends  out  to  occupy 


32  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [382 

the  vacuolated  space  surrounding  the  cirrus  pouch.  The  fully  devel- 
oped uterine  embryos  are  14.5/x  to  15.5/i  in  diameter.  The  inner  embry- 
onic membrane  is  18/*  to  20/*  across;  the  length  of  the  embryo  plus  the 
pyriform  body  is  26.5/*  to  29/*.  The  outer  mebrane  is  55  to  61/*  in 
diameter;  the  middle  is  loosefitting  and  irregular. 

The  ventral  excretory  duct  has  a  diameter  of  45  to  75/*,  and  the 
dorsal  is  about  half  this  size.  Both  lie  in  approximately  the  same 
dorsoventral  plane.  Near  the  distal  end  of  the  proglottid  the  ventral 
duct  turns  and  runs  abruptly  mediad  for  a  distance  of  about  275/* 
where  it  gives  off  the  transverse  commissure.  Beyond  the  commissure 
and  therefore  in  the  next  proglottid  the  duct  turns  laterad  not  so 
abruptly  as  before  to  regain  its  lateral  position.  Stiles  records  that  he 
has  observed  the  dorsal  duct  in  open  communication  with  the  trans- 
verse commissure;  since  this  occurs  also  in  8.  variabilis  and  8.  anoplo- 
cephaloides,  it  seems  to  be  a  common  occurrence  in  the  genus. 

Stiles  in  1896  referred  to  this  species  five  cestodes  from  a  rabbit 
(host  species  and  locality  not  known),  giving  them  the  rank  of  a  va- 
riety with  the  name  Bertia  americana  leporis.  His  description  is  very 
unsatisfactory  being  based  upon  poorly  preserved,  unsectioned  material. 
In  reading  over  carefully  the  description  of  the  worms  there  does  not 
appear  a  single  anatomical  character  upon  which  to  justify  giving 
them  the  rank  of  a  distinct  variety.  The  cirrus  is  supposed  to  be 
smooth ;  but  this  conclusion  is  based  upon  extruded  cirri,  and  as  shown, 
extruded  cirri  of  the  forms  from  the  porcupines  are  usually  if  not 
always  smooth.  The  testes  are  stated  to  be  probably  fewer;  but  Stiles 
is  not  certain  as  to  the  count  and  the  difference  moreover  is  not  so 
great  as  that  between  the  extremes  of  the  next  species.  The  description 
of  the  position  of  the  genital  pore  is  vague  but  implies  a  difference; 
yet  his  figures  (Stiles  1896,  PI.  X,  Figs.  7,  14,  and  15)  are  identical 
for  both.  Some  slight  differences  were  observed  in  the  first  appearance 
of  the  genital  anlagen ;  but  such  appearances  when  based  upon  unsec- 
tioned specimens  and  poor  material,  do  not  have  any  importance.  No 
other  points  of  difference  were  brought  out  which  could  not  be  ac- 
counted for  as  mere  individual  variations.  There  appears  therefore  to 
be  no  evidence  to  justify  giving  these  specimens  the  rank  of  variety. 

On  the  other  hand  there  is  very  little  evidence  that  these  cestodes 
rightly  belong  to  this  species,  or  even  to  the  genus  for  that  matter. 
As  will  be  shown  in  the  succeeding  paragraphs  the  supposedly  homo- 
geneous materials  from  the  porcupine  prove  to  be  two  distinct  species. 
Until  something  is  known  of  them  however  they  should  not  be  given 
separate  recognition. 


383]  AN0PL0CEPHAL1DJE—D0UTHITT  33 

Schizotaenia  variabilis  sp.  nov. 
[Figures  30  to  32] 

Some  years  ago  Dr.  A.  K.  Fisher  collected  cestodes  from  a  number 
of  porcupines  (Erethizon  dorsatus)  at  Lake  George,  New  York.  These 
in  the  Bureau  of  Animal  Industry  at  present  bear  serial  number  1502. 
Stiles  (1896)  examined  them  and  identified  them  as  Bertia  americana 
(now  Schizotaenia  americana).  Professor  Ward  secured  the  loan  of 
these  specimens.  Examination  has  shown  that  they  do  not  represent 
S.  americana,  but  are  a  distinct  species.  The  following  description  is 
based  upon  a  single  complete  specimen,  not  sectioned,  and  several  sec- 
tioned portions  from  the  region  of  sexual  maturity.  The  specimens 
reported  by  Cohn  (1906)  from  Erethizon  epixanthus  from  Alaska  seem 
referable  to  this  species  rather  than  S.  americana,  since  he  reports  that 
the  female  glands  are  located  far  from  the  median  line. 

The  complete  specimen  at  hand,  which  appears  to  be  fully  grown 
and  has  shed  proglottids,  is  20  mm.  long  and  is  composed  of  60  pro- 
glottids.  The  greatest  width,  8.5  mm.,  is  reached  18  cm.  from  the 
head;  back  of  this  the  strobila  narrows  somewhat.  All  the  proglottids 
are  very  short  in  comparison  with  the  width,  except  the  last  which  is 
6  mm.  wide  and  1.5  mm.  long.  The  fragments  at  hand  agree  with  this 
specimen,  except  that  some  have  longer  and  thicker  proglottids  the 
width  being  the  same ;  this  would  seem  to  indicate  that  they  came  from 
a  larger  specimen.  Three  scolices  are  present;  they  measure  450/t  long 
by  875/i  wide.  One  is  retracted  within  the  end  of  the  strobila  as  in 
S.  americana. 

The  genital  pores  are  regularly  alternate.  The  genital  cloaca  and 
cirrus  pouch  agree  with  the  description  just  given  for  S.  americana, 
except  that  the  cirrus  pouch  is  somewhat  smaller  being  ordinarily  485 
to  500/u,  long  tho  it  is  sometimes  longer.  Its  width  is  140  to  185/t.  The 
vesicula  seminalis  is  of  necessity  much  shorter  than  in  S.  americana  on 
account  of  the  nearness  of  the  female  glands  to  the  lateral  margin. 
It  is  variable  in  form,  being  sometimes  straight  and  sometimes  in  the 
form  of  large,  wide  loops,  as  shown  in  Fig.  30.  The  gland-cells  on  the 
inner  end  of  the  vesicula  are  usually  more  numerous  than  in  8.  ameri- 
cana, as  shown  in  the  same  figure.  The  vagina  is  identical  but  of 
course  shorter. 

The  testes  are  decidedly  unlike  8.  americana  in  form  and  distri- 
bution. Nearly  all  are  elongated  anteroposterior^,  the  breadth  being 
to  the  length  as  5 :7.  In  some  specimens  they  are  60  to  80/x,  long  while 
in  others  they  are  95  to  130/x  long.  These  measurements  were  all  taken 
at   sexual  maturity.     They  occur  only  in  the  median  field  and  are 


34  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [384 

mostly  on  the  pore  side  of  the  median  line  of  the  proglottid.  Beyond 
the  median  line  they  are  reduced  to  a  single  row  which  reaches  as  far 
as  the  median  axis  of  the  female  glands  of  the  proglottids  before  and 
behind.  On  the  pore  side  of  the  field  they  are  in  2y2  to  5  rows  when 
viewed  from  above.  They  are  usually  limited  to  the  dorsal  part  of  the 
proglottid  but  in  one  specimen  they  sometimes  extend  entirely  through 
the  medullary  portion.  Generally  they  lie  almost  entirely  posterior  to 
the  female  glands,  but  in  one  specimen  they  were  found  sometimes  at 
the  same  level  as  the  female  glands  and  dorsal  to  them,  and  in  some 
proglottids  even  anterior  to  them,  and  therefore  if  judged  by  position 
in  the  proglottid  in  front!  These  different  types  of  distribution  were 
all  found  in  five  adjacent  proglottids.  The  number  of  testes  was  found 
to  vary  from  60  to  70  in  one  specimen  to  110  in  another.  The  number 
seems  to  be  fairly  constant  for  any  given  individual. 

As  already  stated  the  genital  organs  alternate  regularly  from  the 
left  to  the  right  side  of  the  strobila.  The  median  line  of  the  female 
glands  in  mature  proglottids  is  600/t  from  the  median  line  of  the  pro- 
glottid ;  in  S.  americana  the  distance  was  55  to  175/t.  The  ovary  is  much 
smaller  than  in  S.  americana,  its  diameter  varying  from  650  to  900/x; 
it  never  reaches  to  the  median  line  of  the  proglottid.  Otherwise  the 
ovary  is  not  different  from  that  of  S.  americana.  The  ova  measure  12 
to  14/x  in  diameter.  The  vitelline  gland  shows  the  peculiar  mulberry- 
like appearance  figured  by  von  Janicki  for  S.  hagmanni.  In  cross 
sections  of  the  gland,  i.  e.,  longitudinal  sections  of  the  proglottid,  about 
25  radiating  lobes  can  be  made  out.  There  is  no  trace  of  the  usual 
bilobed  condition.  The  shell  gland  is  small  and  lies  directly  dorsad 
of  the  vitelline  gland,  having  about  one-third  the  diameter  of  the  latter. 
The  uterus  in  mature  proglottids  is  apparently  not  so  extensive  as  in 
8.  americana,  but  statements  concerning  it  must  be  qualified  on  account 
of  the  poor  condition  of  the  material.  It  does  not  seem  to  reach  either 
the  anterior  or  the  posterior  border  of  the  segment.  Otherwise,  the 
description  given  for  8.  americana  will  apply  equally  well  for  the  early 
stages.    Unfortunately  the  later  development  could  not  be  followed. 

The  description  given  for  the  excretory  system  of  S.  americana 
will  answer  for  what  seems  to  be  the  normal  condition  here.  One 
individual  however  offers  such  radical  departures  from  this  type  that 
a  detailed  description  is  given  here.  In  the  specimen  in  question  the 
longitudinal  portions  of  both  dorsal  and  ventral  ducts  are  of  approxi- 
mately the  same  size.  In  one  sectioned  portion  having  five  pro- 
glottids, the  large  ventral  transverse  commissure  extends  laterad  to 
make  open  connection  also  with  the  dorsal  duct.  Judged  by  its  appear- 
ance, the  commissure  would  seem  to  belong  to  the  dorsal  rather  than 


385]  ANOPLOCEPHALIDJE—DOUTHITT  35 

the  ventral  system  in  these  cases,  and  the  connection  with  the  ventral 
system  appears  secondary.  This  effect  is  heightened  by  another  re- 
markable feature,  the  presence  of  a  secondary  transverse  commissure 
anterior  to  the  first.  This  duct  is  a  direct  continuation  mediad  of  the 
longitudinal  ventral  duct  at  the  point  where  the  latter  dips  distad  just 
before  communicating  with  the  main  transverse  commissure.  This  sec- 
ondary duct  is  usually  of  small  diameter  dorsoventrally  but  antero- 
posterior^ it  may  extend  through  one-fourth  the  length  of  the  pro- 
glottid. Its  diameter  however  is  very  irregular,  and  its  course  is  sinu- 
ous. These  secondary  commissures  are  apparently  analogous  to  the 
condition  found  in  8.  anoplocephaloides.  The  connections  between  dor- 
sal ducts  and  transverse  commissures  were  observed  also  in  S.  anoplo- 
cephaloides, and  by  Stiles  in  S.  americana.  In  two  proglottids  the  main 
transverse  commissures  are  not  fully  developed,  but  are  represented  by 
strings  of  disconnected  spaces.  This  may  be  due  however  to  the  fact 
that  the  proglottids  are  not  quite  mature.  All  these  conditions  are 
illustrated  in  Fig.  31. 

Schizotaenia  anoplocephaloides  sp.  nov. 
[Figures  33  to  40] 

Four  gophers  (Geomys  breviceps  Baird)  from  creek  bottom  land 
at  Norman,  Oklahoma,  yielded  of  this  species  plentifully.  About  half 
of  the  75  specimens  taken  were  lost  and  tho  many  others  are  not  in  the 
best  condition,  some  are  good.  For  distribution  and  frequency  of  this 
species  see  the  table  on  page  62. 

The  total  length  of  the  specimens  at  hand  is  30  to  33  mm.  The 
number  of  proglottids  varies  from  55  to  80,  the  average  being  68. 
There  is  no  neck.  The  first  proglottids  are  about  seven-tenths  the 
diameter  of  the  scolex  and  from  one-eighth  to  one-twelfth  as  long  as 
broad.  The  maximum  width  of  1.7  to  2  mm.  is  reached  about  the 
middle  of  the  strobila;  back  of  this  the  width  remains  constant  or 
decreases  slightly.  At  the  middle  of  the  strobila  the  proglottids  are 
from  one-fifth  to  one-third  as  long  as  broad.  They  increase  in  length  as 
they  become  older,  becoming  finally  two-thirds  as  long  as  broad.  In 
those  proglottids  in  which  the  eggs  are  being  fertilized  the  large 
everted  genital  pore  protrudes  prominently. 

The  scolex  averages  390/x  in  diameter  and  320/t  long  and  is  quite 
distinct  from  the  strobila.  It  is  clearly  separated  into  anterior  and 
posterior  portions  by  a  slight  transverse  constriction  near  its  middle. 
The  apex  is  blunt,  the  anterior  outline  being  a  regular  curve.  The 
scolex  is  rather  deeply  four-lobed  owing  to  the  presence  of  longitudinal 


36  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [386 

grooves  between  the  suckers,  the  dorsal  and  ventral  grooves  being  the 
deepest.  All  four  fade  out  before  reaching  the  apex.  The  very  promi- 
nent suckers  face  anterolaterad  at  an  angle  of  about  45  degrees  from 
the  axis  of  the  worm.  The  mouths  of  the  suckers  are  from  25  to  40/u, 
in  diameter  and  the  cavities  are  70  to  90 ■/*  deep.  The  muscular  wall  of 
the  sucker  is  from  25  to  50/a  thick. 

There  is  one  set  of  reproductive  organs  to  the  proglottid,  these 
alternating  with  practically  perfect  regularity  from  right  to  left;  in  a 
dozen  worms  examined  only  three  cases  were  found  of  two  adjacent 
proglottids  with  pores  on  the  same  side.  The  anlagen  of  the  female 
glands  can  be  made  out  in  the  first  proglottid.  Testes  are  to  be  recog- 
nized in  the  15th  and  eggs  pass  into  the  uterus  in  the  40th.  The 
genital  cloaca  presents  an  unusual  appearance.  It  is  clearly  divisible 
into  two  parts,  a  lateral  and  a  median.  The  lateral  is  an  unusually 
large  structure  varying  in  shape  in  different  specimens  probably  on 
account  of  contraction  from  nearly  globular  to  angular,  being  even 
acutely  angular  at  its  inner  end.  The  pouch  has  a  length  of  about  140/t 
and  a  breadth  of  about  85/*.  The  anlage  of  this  structure  can  be  made 
out  in  about  the  20th  proglottid  as  a  group  of  cells  that  stain  deeply. 
Soon  afterwards  a  mass  of  substance  not  distinguishable  from  the  cuti- 
cula  can  be  made  out  in  it  near  the  lateral  margin.  A  lumen  appears 
which  increases  in  length  mediad  and  then  expands.  Just  before  sexual 
maturity  the  cuticula  of  the  outer  body  wall  is  broken  through,  and  the 
cavity  of  the  pocket  becomes  continuous  with  the  outer  surface.  The 
pouch  immediately  turns  inside  out,  projecting  abruptly  in  some  cases 
for  more  than  '200/i  from  the  margin.  Fifteen  or  twenty  proglottids 
farther  back  it  is  again  retraced  to  remain  thus  to  the  end.  In  its 
fully  developed  stage  the  cuticula  and  wall  of  the  pocket  are  directly 
continuous  with  the  cortical  layer  and  not  distinguishable  from  it. 
Mediad  of  this  and  communicating  with  it  is  a  second  small  pocket. 
It  is  rather  small  and  is  expanded  at  its  inner  end  to  embrace  the  end 
of  the  cirrus  pouch.  The  vagina  opens  into  its  anterior  border.  Gen- 
erally, the  median  end  of  the  cloaca  lies  mediad  of  the  longitudinal 
excretory  ducts. 

An  unexpected  condition  was  found  in  the  position  of  the  vagina 
and  cirrus  pouch  with  respect  to  each  other  and  to  the  nerve  and 
excretory  trunks.  When  the  pouch  and  vagina  are  on  the  right  margin 
of  the  strobila  they  cross  the  excretory  ducts  dorsally ;  but  when  on  the 
left,  in  two  individuals  out  of  ten  studied,  they  more  often  cross  the 
excretory  ducts  ventrally.  In  these  two  individuals  out  of  19  proglot- 
tids which  had  the  pores  on  the  left  margin  ten  had  the  genital  ducts 
ventral.     Thus  some  proglottids  have  an  arrangement  that  is  charac- 


387]  ANOPLOCEPHALIDJE—DOUTHITT  37 

teristic  of  the  Linstowinae  and  has  been  observed  nowhere  amongst  the 
Anoplocephalinae,  except  possibly  in  the  genus  Triplotaenia.  It  should 
be  stated  however  that  the  identification  of  right  and  left  sides  in  these 
specimens  may  be  in  error. 

Still  another  interesting  feature  was  observed  in  regard  to  the 
genital  ducts.  The  vagina  crosses  the  cirrus  pouch  ventrally  in  most 
cases;  but  in  the  two  individuals  mentioned  in  the  paragraph  above,  12 
of  the  proglottids  that  had  the  excretory  ducts  dorsal  to  the  genital 
ducts,  had  the  vagina  dorsal  and  the  cirrus  pouch  ventral.  This  recalls 
at  once  Moniezia  in  which  the  vagina  is  regularly  ventral  to  the  cirrus 
pouch  on  the  right  and  dorsal  on  the  left.  These  same  two  individuals 
were  unusual  in  the  form  of  the  ovary,  as  explained  below.  The  cirrus 
pouch  is  very  large.  Before  the  evagination  of  the  cloaca  it  lies  usually 
entirely  mediad  of  the  excretory  ducts;  but  when  the  cloaca  is  everted 
it  may  lie  entirely  laterad.  In  mature  proglottids  it  is  275  to  370/x 
long,  and  120  to  240/x  broad.  Before  the  development  of  the  vesicula 
seminalis  it  may  be  elongated  and  spindleshaped ;  afterwards  it  is  al- 
ways pearshaped.  The  cirrus  is  straight  in  the  lateral  half  of  its 
length  and  has  usually  two  spiral  turns  at  its  upper  end.  It  is  lined 
with  many  hundreds  of  small  hooks  which  are  arranged  in  spiral  rows. 
The  median  end  of  the  cirrus  pouch  is  occupied  by  a  portion  of  the 
vesicula  seminalis.  Proximad  and  mediad  of  the  pouch  is  a  second 
portion,  140/x  long,  which  is  only  slightly  coiled.  This  portion  is  thickly 
beset  with  glandular  cells.  Beyond  the  seminal  vesicle  the  passage 
narrows  and  soon  divides,  sending  a  branch  to  either  testicular  field. 

The  testes,  70  to  110  in  number,  are  dorsal  and  posterior.  They 
occur  in  both  lateral  halves  but  are  mostly  on  the  side  away  from  the 
pore.  At  either  side  they  are  grouped  two  or  three  deep  dorsoven- 
trally.  Above  the  yolk-gland  there  is  never  more  than  a  single  layer 
and  as  often  as  not  there  are  none  here,  the  testes  being  separated 
into  two  groups.  The  testes  are  typically  spherical  in  shape,  averaging 
35/t  in  diameter. 

The  vagina  communicates  with  the  genital  cloaca  on  the  anterior 
surface  of  the  latter.  It  extends  forward  and  somewhat  mediad  in 
front  of  the  cirrus  pouch  then  turns  directly  toward  the  median  line, 
usually  crossing  the  cirrus  pouch  ventrally.  Mediad  of  the  cirrus  pouch 
it  curves  somewhat  distad,  the  entire  vagina  thus  describing  a  semi- 
circle. Its  middle  portion  in  the  anterior  part  of  the  proglottid  is 
several  times  the  diameter  of  the  two  ends.  The  surface  is  glandular. 
At  about  the  same  level  as  the  lateral  end  is  located  the  receptaculum 
seminis  which  is  a  simple  expansion  of  the  vagina,  measuring  180  by 


38  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [388 

130/*.  The  ovary  is  distinctly,  but  not  considerably,  to  the  pore  side  of 
the  median  line.  It  is  fanshaped  in  outline,  being  formed  of  a  large 
central  mass  from  which  about  30  lobes  radiate  extending  in  all  direc- 
tions in  the  horizontal  plane  except  towards  the  part  of  the  field  occu- 
pied by  the  vitelline  gland.  It  is  distinctly  ventral  in  position.  In 
two  specimens  however  most  of  the  proglottids  which  have  the  genital 
pore  on  the  left  side  have  the  ovary  separated  into  distinct  dorsal  and 
ventral  portions  with  a  narrow  connection  between  them.  The  dorsal 
portion  in  these  cases  is  of  about  half  the  area  of  the  ventral  and  thin. 
These  are  the  individuals  that  show  the  peculiarities  of  the  excretory 
ducts  already  mentioned.  The  oviduct  in  all  cases  connects  with  the 
ovary  at  a  point  directly  in  front  of  the  vitelline  gland  and  is  short. 
The  vitelline  gland  is  of  the  ordinary  bilobed  type  which  is  found  nearly 
everywhere  in  the  subfamily  and  shows  no  suggestion  of  the  type  found 
usually  in  the  genus.  It  lies  posteriorly  to  the  ovary,  sometimes  ex- 
tending dorsad  to  the  cortical  layer  and  dividing  the  testes  into  two 
fields.  In  cross  section  it  is  distinctly  horseshoeshaped,  with  the  open 
face  dorsad.  Viewed  from  above  it  is  U-shaped  but  with  the  median 
arm  much  larger. 

The  uterus  is  first  recognizable  as  a  sheet  of  deeply  staining  cells 
ventrad  of  the  testes  and  dorsad  of  the  ovary.  Just  beneath  the  mar- 
gin of  each  testicular  field  the  sheet  is  thickened  into  a  heavy  circular 
band.  On  the  pore  side  there  extends  forward  from  the  anterior  tip 
of  the  circular  band  a  strip  of  the  same  nature,  which  crosses  the  vagina 
laterad  of  the  receptaculum  seminis  then  turns  and  extends  diagonally 
distad  just  underneath  the  anterior  margin  of  the  receptaculum  and 
anterior  to  the  vitelline  gland.  It  joins  with  the  other  circular  band  at 
the  corner  of  this  gland.  It  is  into  this  cross  duct  that  the  uterine  duct 
empties.  Within  these  circles  and  anterior  to  the  transverse  portion 
and  connecting  with  them  at  frequent  points,  is  a  network  of  strands 
formed  by  thickenings  of  the  uterine  tissue  which  recalls  strongly  the 
reticulate  uterus  found  in  Moniezia.  The  transverse  portion  is  of  course 
the  first  part  of  the  uterus  to  receive  eggs;  but  the  eggs  pass  immedi- 
ately into  the  circular  bands  which  have  developed  into  canals.  The 
extreme  lateral  portions  fill  and  then  the  whole  canals.  As  the  uterus 
fills  with  eggs  these  passages  expand  centrally  and  the  transverse  por- 
tion anteriorly,  so  that  the  cavity  of  the  uterus  becomes  one  continuous 
sac.  The  extension  from  this  stage  is  by  regular  outpocketing  no 
different  from  that  found  in  Anoplocephala  and  Bertiella,  except  that 
the  pockets  are  of  necessity  shorter  since  the  uterus  does  not  begin  as  a 
simple  tube.  The  development  of  the  uterus  continues  until  nearly  all 
the  organs  of  the  proglottid  are  reabsorbed.     The  cirrus  pouch  and 


389]  AN0PL0CEPHALID2E—D0UTHITT  39 

vesicula  persist  to  the  last  either  in  situ  or  crowded  to  one  side.  The 
receptaeulum  persists  as  a  mere  vestige.  The  shell  gland  persists  as  a 
structureless  mass  connected  with  the  receptaeulum  and  with  the  pos- 
terior end  of  the  proglottid  by  a  narrow  bridge,  such  as  has  been  de- 
scribed for  Moniezia  carrinoi  and  some  of  its  relatives.  All  the  other 
organs  are  reabsorbed,  the  testes  being  the  last  to  give  way.  The  pro- 
glottid becomes  little  more  than  a  thin-walled  egg  capsule,  the  cortical 
layer  being  reduced  sometimes  to  a  thickness  of  20//.. 

The  uteri  of  the  three  species  of  Schizotaenia  here  described  agree 
in  all  essential  respects  but  differ  decidedly  from  the  account  presented 
by  von  Janicki  (1906)  for  S.  hagmanni  which  is  accepted  as  the  type 
of  the  genus.  He  recognized  nothing  comparable  to  the  degenerate 
reticulum  or  diffuse  uterus  described  here;  also  the  three  species  de- 
scribed here  show  nothing  comparable  to  the  "ausserst  feinen  Schlitze, 
der  als  mannigfach  gewellte  Flache  durch  den  grossten  Teil  der  Mark- 
schicht  *****  sich  hinzieht".  Von  Janicki  does  not  figure  this 
structure  except  in  transverse  section,  in  which  way  it  gives  very  little 
information.  In  view  however  of  the  fact  that  such  a  structure  would 
not  be  greatly  different  in  appearance  from  that  of  the  degenerate  retic- 
ulum described  here,  I  am  inclined  to  believe  the  difference  is  one  of 
interpretation  rather  than  structure.  Likewise  von  Janicki 's  drawings 
of  the  ' ' Spaltenwerk "  (Figs.  85,  86)  resemble  closely  the  condition 
observed  in  frontal  sections  of  the  early  saccular  stage  when  the  thin 
uterus  is  applied  to  the  uneven  surface  of  the  ovary  and  only  parts 
of  the  cavity  appear  in  each  section.  There  is  no  ground  therefore 
for  considering  these  uteri  as  differing  fundamentally  in  this  regard. 
The  possession  of  uterine  outpocketings  by  the  present  species  is  strik- 
ing and  is  significant  as  to  its  relationships;  but  the  fact  offers  no 
barrier  to  its  inclusion  in  the  genus  Schizotaenia;  it  is  what  should  be 
expected  in  the  more  primitive  members  of  the  genus. 

The  uterine  embryos  have  three  membranes.  The  outer  is  spherical 
and  30  to  40//.  in  diameter.  The  middle  is  loose-fitting  and  irregular. 
The  diameter  of  the  embryo  is  10  to  17//  with  no  considerable  range  of 
variation  in  any  one  individual.  There  is  a  perfectly  developed  pyri- 
form  apparatus  whose  length  plus  that  of  the  embryo  is  16  to  26/x.  The 
two  stout  tapering  horns  cross  at  the  tip  and  extend  backwards  as 
long,  slender  processes. 

The  excretory  ducts  lies  in  about  the  same  dorsoventral  plane,  the 
originally  dorsal  duct  outside.  This  arrangement  is  present  in  both 
strobila  and  scolex.  The  path  across  the  proglottid  is  a  curve.  The 
dorsal  duct  is  usually  between  10  and  12//,  in  diameter  at  the  ends  of 
the  proglottids,  narrowing  in  the  middle  to  6  or  7//,  on  the  side  away 


40  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [390 

from  the  pore,  and  on  the  pore  side  to  3/*.  The  ventral  duct  at  the 
ends  of  the  proglottids  is  40  to  50/*  in  diameter ;  it  narrows  in  the  middle 
to  30  to  35fi  on  the  side  away  from  the  pore,  and  on  the  pore  side  to 
3  by  10ft.  The  transverse  commissure  of  the  ventral  duct  is  a  straight 
tube  25  to  40ft  in  diameter.  In  three  individuals  the  dorsal  longitu- 
dinal duct  is  connected  with  the  transverse  commissure  in  very  many 
cases  (Fig.  40).  In  the  present  species  these  connections  are  always 
on  the  same  side  of  the  strobila,  judged  to  be  the  right  side,  tho  since 
the  specimens  are  in  longitudinal  sections  this  could  not  be  determined 
finally. 

In  the  anterior  aporose  portion  of  the  proglottid  the  transverse 
commissure  becomes  divided  and  with  the  aid  of  the  main  duct  sur- 
rounds an  "island"  of  medullary  parenchyma  which  varies  from  60 
to  200ft  long  and  may  have  its  long  axis  either  longitudinal  or  trans- 
verse. From  this  region  but  varying  considerably  in  point  of  origin 
a  large  branch  duct  extends  distad  and  mediad  into  the  proglottid. 
This  duct  persists  after  the  uterus  is  fully  formed,  projecting  free  into 
its  cavity  and  invested  by  its  walls.  Frequently  a  similar  duct  will 
extend  forward  from  the  transverse  commissure  in  the  lower  pore  corner 
of  the  proglottid. 

There  appears  to  be  no  doubt  but  that  this  cestode  is  most  nearly 
allied  to  the  genus  Schizotaenia.  Yet  it  shows  very  clearly  that  it  is 
related  to  the  genus  Anoplocephala  and  appears  to  be  transitional  be- 
tween the  two.  Aside  from  the  close  general  resemblances  between  the 
two  genera  the  following  characters  show  its  close  relationship  to  the 
known  representatives  of  the  genus  Anoplocephala. 

1.  The  testes  are  mainly  on  the  side  away  from  the  pore. 

2.  The  vitelline  gland  is  of  the  type  found  in  Anoplocephala,  and 
shows  no  suggestion  of  the  type  occurring  in  most  other  known  Schizo- 
taenia. 

3.  The  uterus  develops  by  regular  anterior  and  posterior  out- 
pocketing. 

4.  The  cirrus  pouch  is  unusually  large. 

THE  GENUS  SCHIZOTAENIA 

Deiner  (1912)  has  published  an  account  of  the  anatomy  of  Taenia 
magna  Murie  1870,  under  the  name  Anoplocephala  latissima,  the  specific 
name  "magna"  being  dropped  because  preoccupied  by  A.  magna  Abild- 
gaard  1789.  The  account  is  very  thoro  and  bears  all  the  marks  of  being 
strictly  dependable.  A  consideration  of  the  anatomical  features  leaves 
no  doubt  whatever  in  my  mind  but  that  this  cestode  is  a  true  repre- 


391]  ANOPLOCEPHALIDJE—DOUTHITT  41 

sentative  of  the  genus  Schizotaenia.    Briefly  summarized,  the  following 
points  brought  out  by  Deiner  support  this  conclusion. 

1.  The  testes  are  mainly  on  the  pore  side;  this  agrees  with  some 
Schizotaeniae  and  disagrees  with  all  known  Anoplocephalae. 

2.  The  cirrus  is  spiny. 

3.  The  vagina  and  vaginal  pore  are  anterior  to  the  cirrus  pouch. 

4.  The  structure  of  the  vagina  is  identical  with  that  of  8.  anoplo- 
cephaloides. 

5.  The  oviduct  connects  with  the  ovary  directly  in  front  of  the 
vitelline  gland. 

6.  The  uterus,  while  relatively  simple,  shows  clearly  a  resemblance 
to  Schizotaenia. 

These  features  are  all  characteristic  of  the  representatives  of  the 
genus  Schizotaenia.  Some  of  them  are  found  nowhere  else;  none  of 
them  are  found  in  any  species  known  to  belong  to  the  genus  Anoplo- 
cephala.  In  one  character  only  does  this  cestode  resemble  Anoplo- 
cephala:  the  genital  pores  are  all  dextral.  This  character  however  can 
not  be  considered  to  outweigh  the  other  anatomical  resemblances  and 
the  cestode  is  therefore  placed  in  the  genus  Schizotaenia. 

MacCallum  and  MacCallum  (1912)  have  published  a  careful  ac- 
count of  the  anatomy  of  Taenia  gigantea  Peters  1856,  usually  referred 
to  the  genus  Anoplocephala.  This  cestode  also  shows  marked  affinities 
for  Schizotaenia  and  while  the  case  is  not  so  strong  as  in  the  other, 
the  evidence  is  sufficient  to  indicate  the  position  of  the  cestode.  The 
evidence  that  Taenia  gigantea  Peters  is  a  Schizotaenia  follows. 

1.  The  testes  are  mostly  on  the  pore  side. 

2.  The  cirrus  is  spiny. 

3.  The  vagina  was  not  observed.  Since  the  investigators  expected 
to  find  it  posterior  to  the  cirrus  pouch,  an  inconspicuous  and  transitory 
vagina,  such  as  is  found  in  8.  americana  and  8.  variabilis,  anterior  to 
the  pouch  might  easily  have  been  overlooked. 

4.  The  vitelline  gland  is  lobulated. 

Taenia  gigantea  Peters  is  therefore  here  transferred  to  the  genus 
Schizotaenia.  Its  specific  distinctness  from  8.  magna,  long  disputed, 
seems  assured. 

With  six  of  the  species  of  Shizotaenia  well  known  it  is  now  possible 
to  judge  the  value  of  the  various  anatomical  features  much  more  satis- 
factorily than  could  von  Janicki,  who  had  before  him  but  one  that  was 
satisfactorily  described.  It  becomes  evident  that  several  of  the  charac- 
teristics he  proposed  must  be  restated  and  several  others  show  them- 
selves to  be  of  generic  rank.    In  making  such  generalizations  the  poorly 


42  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [392 

known  S.  decrescens  should  not  be  given  serious  consideration.  Its 
position  in  the  genus  is  not  assured  and  the  statements  concerning  it  are 
not  clear-cut  and  dependable.  The  conception  of  the  genus  must  be 
changed  in  the  following  particulars: 

1.  The  testes  do  not  necessarily  extend  "von  Langsgefass  zu 
Langsgefass"  in  the  distal  part  of  the  proglottid.  They  may  be  in  two 
lateral  groups,  or  all  on  the  pore  side,  and  they  may  be  mostly  anterior. 

2.  In  the  main  the  genital  canals  "ziehen  dorsal  an  den  beiden 
Excretionsgefassen  und  dem  Nervenstrang  vorbei";  yet  recognition 
must  be  given  to  the  fact  that  the  opposite  may  be  the  case  in  some 
proglottids. 

3.  The  position  of  the  vagina  and  vaginal  pore  anterior  to  the 
cirrus  pouch  is  an  absolute  distinction  from  all  other  known  genera  of 
the  subfamily  and  indeed  from  nearly  all  known  cestodes,  and  should 
have  recognition  as  perhaps  the  most  important  generic  character.  The 
unsatisfactory  description  of  S.  decrescens  is  noncommittal  but  gives 
no  information  opposed  to  the  conclusion  that  the  vagina  is  anterior 
here  also. 

4.  The  union  of  the  oviduct  with  the  ovary  in  all  sufficiently 
known  species  of  the  genus  is  located  directly  anterior  to  the  middle 
of  the  vitelline  gland.    This  fact  should  be  stated  in  the  diagnosis. 

5.  The  diagnosis  should  state  that  cirrus  is  spiny. 

Accordingly  the  genus  Schizotaenia  should  be  characterized  as 
follows : 

Anoplocephalinae,  with  segments  broader  than  long.  Genital  pores 
regularly  alternate  or  dextral,  and  in  one  doubtful  species  irregularly 
alternate.  Dorsal  excretory  duct  lateral  of  ventral.  Genital  canals 
pass  usually  dorsal  of  longitudinal  excretory  vessels  and  nerve,  tho  the 
reverse  condition  has  been  observed.  Testes  confined  to  the  median 
field,  either  distal  in  position  or  proximal,  and  mostly  on  the  pore  side. 
Cirrus  pouch  very  large  and  muscular,  cirrus  spiny.  External  vesicula 
seminal  is  present.  Vagina  and  vaginal  pore  anterior  to  the  cirrus 
pouch.  Female  glands  placed  towards  the  pore  side  of  the  median 
field.  Oviduct  joins  the  ovary  directly  in  front  of  the  middle  of  the 
vitelline  gland.  Uterus  not  a  simple  transverse  tube,  usually  perhaps 
always  a  degenerate  reticulum;  confined  to  the  median  field  in  anlage, 
and  in  its  fully  developed  stage  either  there  or  crossing  the  excretory 
ducts  mostly  on  the  dorsal  side.  No  pyriform  apparatus.  Adults  in 
mammals. 

Designated  as  type :    Schizotaenia  decrescens  Diesing  1856. 


393]  ANOPLOCEPHALIDAz— DOUTHITT  43 

The  genus  Schizotaenia  embraces   the   following    well    described 

species. 

Magna  group:    Genital  pores  dextral.    Testes  mostly  anterior  and 

mostly  on  the  pore  side.    Size  of  known  species  enormous. 
S.  latissima  Deiner  8.  gigantea  Peters 

Hagmanni    group:     Genital    pores    alternate.     Testes    posterior. 

Known  species  small. 

S.  hagmanni  von  Janicki  8.  variabilis  Douthitt 

S.  americana  Stiles  S.  anoplocephaloides  Douthitt 

"Schizotaenia"  deerescens  (Diesing  1856)   Liihe  1895,  which  von 

Janicki  unfortunately  designated  as  the  type  of  the  genus,  seems  to 

belong  to  this  group  and  should  be  treated  as  such  until  better  known. 

1 '  Anoplocephala' '  transversaria  shows  some  of  the  characteristics  of  this 

group  but  not  enough  information  is  accessible  to  justify  a  change  of 

position. 

THE  UTERUS  OP  MONIEZIA  EXPANSA 

The  cestodes  of  the  genus  Moniezia  have  received  more  attention 
than  those  of  any  other  genus  of  the  family.  As  a  result  of  the  studies 
of  Stiles  and  Hassall,  Tower,  Fuhrmann,  and  a  number  of  other  work- 
ers, most  of  the  points  of  anatomy  have  been  determined  carefully. 
Strangely  enough,  however,  practically  no  attention  has  been  given  to 
the  uterus  except  to  observe  that  it  begins  usually  as  a  reticulum;  one 
is  left  to  assume  that  the  later  development  is  not  different  from  that 
of  related  forms.  My  own  observations  however  have  tended  to  show 
that  the  development  of  the  uterus  is  in  many  ways  unique  and  fur- 
nishes hints  as  to  the  relationship  of  the  genus.  I  had  for  study  several 
specimens  of  Moniezia  expansa,  taken  from  sheep  at  Lincoln,  Nebraska, 
and  belonging  to  the  collection  of  Professor  Henry  B.  "Ward.  No  ob- 
servations at  variance  with  accepted  anatomical  accounts  were  made 
so  structures  other  than  the  uterus  have  not  been  taken  up,  except  that 
a  drawing  of  a  mature  proglottid  is  included  for  comparison  (Fig.  41). 

The  uterus  in  Moniezia  expansa  begins  as  a  reticulum  (Fig.  42), 
whose  slender  branches  are  quite  distinct  and  lie  at  a  considerable 
distance  from  each  other.  In  extends  nearly  the  whole  length  of  the 
proglottid  in  the  space  that  lies  between  the  shell  glands.  Beyond 
these  glands  on  either  side  two  branches,  occurring  respectively  in  the 
anterior  one-third  and  the  posterior  one-third  of  the  proglottid,  extend 
laterad  to  beyond  the  excretory  ducts  on  the  dorsal  side  of  the  latter. 
Upon  the  entrance  of  the  eggs  the  cavities  of  the  various  branches  of 
the  uterus  enlarge  somewhat.  The  eggs  come  to  lie  in  bunches  which 
are  distributed  equally  to  different  parts  of  the  proglottid.     As  the 


44  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [394 

embryos  develop  the  portions  of  the  uterus  containing  these  bunches 
expand.  This  expansion  continues  until  the  enlarged  spaces  fill  almost 
the  entire  medullary  portion,  the  tissue  substance  beween  the  different 
regions  of  enlargement  being  reduced  greatly  in  thickness  but  remain- 
ing to  the  last  to  separate  these  chambers  from  each  other  (Fig.  43). 
In  no  stage  does  the  uterus  force  its  way  between  the  excretory  ducts 
and  the  ventral  cortical  layer.  Thus  here  in  the  ripe  uterus  is  a  con- 
dition that  is  very  unusual  in  the  Anoplocephalinae,  and  which  indeed 
has  been  observed  in  only  one  other  species  of  the  group,  namely, 
Schizotaenia  americana.  One  recalls  at  once  the  uterus  of  the  Lin- 
stowinae,  tho  there  is  no  indication  that  the  uterus  ever  really  breaks 
down  in  the  present  species.  It  appears  therefore  it  would  not  be  cor- 
rect to  speak  of  the  compartments  as  egg-capsules  in  the  sense  that  the 
term  is  used,  even  tho  it  is  probably  a  step  in  that  direction. 

Drawings  by  Stiles  and  Hassall  (1893)  of  gravid  uteri  of  Moniezia 
pianissimo,  and  M.  alba  indicate  structures  which,  tho  not  so  repre- 
sented, may  be  identical  with  the  walls  of  the  compartments  found  in 
M.  expansa.  There  is  a  slight  indication  of  such  structures  also  in  their 
drawing  of  the  gravid  uterus  of  M.  neumanni.  Their  drawing  of  M. 
trigonophora  shows  no  such  structures  but  since  they  give  no  indica- 
tions of  such  structures  in  their  drawing  of  M .  expansa,  the  absence 
means  nothing.  Thus  no  evidence  available  is  opposed  to  the  conclusion 
that  the  Monieziae  from  mammals  have  the  uterus  similar  in  all  essen- 
tial respects  to  the  condition  here  described  for  Moniezia  expansa.  The 
excellent  account  of  Fuhrmann  (1902)  shows  however  that  the  uteri 
of  the  Monieziae  from  birds  differ  in  almost  every  point  from  the  type 
just  described.  The  difference  between  these  two  groups,  if  the  condi- 
tion for  M.  expansa  is  likewise  the  condition  for  other  mamalian  Monie- 
ziae, may  be  summed  up  as  follows : 

Monieziae  from  mammals  Monieziae  from  birds 

Mature  uterus  crossing  excretory  Mature  uterus   not  crossing  excre- 

ducts  tory  ducts 

Mature  uterus  a  complicated  Mature  uterus  a  simple  transverse 

reticulum  tube,     or     with     a     few     simple 

branches 

Gravid  uterus  in  the  form  of  many  Gravid    uterus    saccular,    or    with 

separated  or  nearly  separted  simple     anterior     and     posterior 

compartments  pockets 

Embryos  with  pyriform  apparatus,  Embryos  without  pyriform  appara- 

the  horns  of  which  end  in  disk  tus 

Dorsal  excretory  duct  median  to  Dorsal  excretory  duct  dorsal  to 

ventral  ventral 


395]  ANOPLOCEPHALIDJE—DOUTHITT  45 

Diamare  (1900)  who  was  first  to  give  more  than  mention  of  the 
Monieziae  from  birds,  created  the  genus  Paronia  for  the  species  he  was 
considering  (Moniezia  carrinoi).  His  conclusions,  however,  were  not 
based  upon  a  sufficient  knowledge  of  the  cestodes  in  question  or  the 
Anoplocephalidae  in  general,  and  Fuhrmann  very  properly  included 
this  cestode  and  other  related  ones  which  he  described  in  the  genus 
Moniezia.  If,  however,  the  additions  here  made  to  the  knowledge  of 
Moniezia  expansa  should  hold  true  for  the  other  Monieziae  from  mam- 
mals, there  seems  to  be  little  question  but  that  the  genus  Paronia  should 
be  revived  for  the  forms  from  birds.  It  would  be  rash  to  conclude, 
however,  that  the  other  Monieziae  from  mammals  agree  with  M .  expansa 
in  this  regard.  If  one  considers  the  different  members  of  the  genus 
Schizotaenia,  one  finds  as  great  a  diversity  in  uterine  structure  as  is  here 
exhibited.  As  to  the  other  differences  noted,  they  are  not  by  themselves 
sufficient  reasons  for  the  separation  of  the  genus  Moniezia,  since  other 
Anoplocephalid  genera  show  both  conditions  for  each  structure.  Any 
division  of  the  genus  Moniezia  would  seem  premature,  therefore,  with- 
out first  examining  several  species.  Consideration  of  this  step  is  left 
to  some  worker  who  has  access  to  such  material. 

For  reasons  stated  on  page  49  I  have  transferred  Zschokke's 
"Moniezia"  diaphana  to  the  genus  Cittotaenia.  Von  Janicki's  descrip- 
tion of  M.  beauforti  is  not  accessible  to  me;  but  since  it  is  an  avian 
parasite,  it  is  here  considered  as  belonging  in  the  carnnoi-group  I  can 
not  find  evidence  to  support  Parona's  action  in  placing  Taenia  frontina 
(Dujardin  1845)  in  this  group.  Until  something  more  is  known  of  this 
cestode  it  should  by  all  means  be  left  where  it  will  do  no  harm.  The 
genus  Moniezia  as  here  accepted  includes  therefore  the  following  species : 

1.  Carrinoi-group  M.  columbae  Fuhrmann 
M.  carrinoi  Diamere  M.  variabilis  Fuhrmann 
M.  ambigua  Fuhrmann                 M.  beauforti  von  Janicki 

2.  Expansa-groxvp 

M.  expansa  Rudolphi  M.  neumanni  Moniez 

M.  trigonophora  Stiles  and  M.  alba  Perroncita 

Hassall  M.  amphibia  von  Linstow 

M .  oblongiceps  Stiles  and  M.  rugosa  Diesing 

Hassall  M.  f estiva  Rudolphi 
M.  planissima  Stiles  and  Hassall 
M.  benedeni  Moniez 

Apparently  in  all  Monieziae  the  oviduct  connects  with  the  ovary 
directly  in  front  of  the  vitelline  gland  and  the  uterus  crosses  the  excre- 


46  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [396 

tory  ducts  dorsally  or  not  at  all.    These  points  should  be  stated  in  the 
diagnosis  of  the  genus. 

The  diagnosis  therefore  becomes  as  follows: 

Anoplocephalinae,  with  segments  generally  broader  than  long. 
Two  complete  sets  of  reproductive  organs  in  each  segment,  or  with  but 
one  continuous  uterus.  Uterus  generally  reticular,  either  confined  to 
the  median  field  or  crossing  the  excretory  ducts  dorsally  only.  Oviduct 
joins  with  ovary  directly  in  front  of  the  middle  of  the  vitelline  gland. 
Genital  canals  cross  on  dorsal  side  of  longitudinal  excretory  vessels  and 
nerves.  Interproglottidal  glands  often  present.  Vagina  ventral  and 
cirrus  dorsal  on  right  side  of  the  proglottid;  the  reverse  on  the  left 
side.  Pyriform  apparatus  present  or  absent;  when  present,  the  horns 
end  in  a  disk.    Adults  in  mammals  and  birds. 

Type-species :    Manieza  expansa  Rudolphi  1810. 

THE  CITTOTAENIAE  OF  NORTH  AMERICAN  RABBITS 

There  have  been  described  and  named  as  parasites  of  North  Ameri- 
can rabbits  four  Cittotaeniae,  namely,  C.  perplexa,  C.  mosaica,  C.  pecti- 
naia,  and  C.  variabilis.  Of  variabilis  I  had  at  hand  the  material  studied 
by  Lyman  (1902)  and  several  specimens  I  collected  at  various  points 
in  Kansas  and  Oklahoma.  The  observations  confirm  Lyman's  state- 
ments and  need  be  discussed  no  farther.  A  drawing  is  given  (Fig.  44) 
for  purposes  of  comparison. 

Of  C.  pectinata  there  is  at  hand  Lyman's  material  from  Kansas 
and  Nebraska  and  also  two  individuals  which  I  collected  at  Neosho 
Falls,  Kansas.  I  have  been  unable  to  find  any  confirmation  of  Lyman's 
statement  that  the  uterus  in  anlage  extends  beyond  the  excretory  ducts. 
Stiles  (1896)  and  Hall  (1908)  also  conclude  that  it  does  not.  In  some 
instances  I  have  observed  that  the  fully  developed  uterus  extends 
slightly  beyond  the  excretory  ducts  dorsally;  but  usually  it  is  confined 
to  the  median  field  at  all  stages.  Lyman  states  that  the  greatest  diame- 
ter of  the  lobes  or  pouches  of  the  ovary  is  16/*.  My  own  measurements 
of  Lyman's  material  and  my  own  show  a  diameter  of  40  to  60/z  as  the 
usual  condition.  Since  the  ova  themselves  are  15  to  18/x  in  diameter 
his  figures  seem  most  probably  a  typographical  error. 

The  specimens  taken  at  Neosho  Falls,  Kansas,  are  in  some  respects 
quite  different  from  Lyman's.  The  testes  are  practically  absent  from 
the  middle  part  of  the  proglottid  and  somewhat  more  numerous  in  the 
lateral  parts  than  in  Lyman's  material.  These  two  specimens  which 
have  shed  some  proglottids  are  45  mm.  long  and  consist  each  of  85 
proglottids.    Lyman's  specimens  were  up  to  71  mm.  long  and  had  when 


397]  AN0PL0CEPHAL1DJE—D0UTHITT  47 

complete  110  to  140  proglottids;  Stiles'  material  measured  up  to  400/* 
long.  Eggs  begin  to  pass  into  the  uterus  in  the  45th  proglottid  in  one 
of  my  own  specimens.  My  measurements  of  the  heads  of  the  two 
specimens  show  diameters  at  the  base  of  700  and  745/*  respectively  but 
confirm  Lyman's  measurements  for  the  specimens  he  studied.  There 
seems  no  reason  to  doubt  Stiles'  figures  which  give  the  diameter  of  the 
head  as  250/*  or  less.  These  different  figures  show  little  reliance  is  to 
be  placed  upon  these  characters. 

In  certain  features  the  American  representatives  of  this  species 
are  different  from  the  European  and  should  be  designated  as  a  distinct 
variety  to  avoid  confusion.    The  following  is  the  characterization. 

Cittotaenia  pectinata:  Strobila  up  to  400  mm.  long.  Sexual  ma- 
turity about  30  mm.  from  the  anterior  end.  Testes  about  150,  extending 
uniformly  across  the  distal  end  of  the  proglottid,  from  excretory  duct 
to  excretory  duct.  Hosts  Lepus  timidus  and  Lepus  variabilis.  Known 
distribution,  Germany  and  France. 

Cittotaenia  pectinata  americana:  Strobila  44  to  71  mm.  long. 
Number  of  proglottids,  85  to  140.  Sexual  maturity  reached  in  about 
the  45th  proglottid,  about  10  mm.  from  the  anterior  end.  Testes  100 
to  125,  extending  from  excretory  duct  to  excretory  duct  in  the  distal 
end  of  the  proglottid,  sometimes  nearly  absent  from  the  median  part 
of  the  field.  Host,  Lepus  californicus  melanotus.  Known  distribution, 
eastern  Kansas  and  Nebraska,  U.  S.  A. 

Of  Cittotaenia  perplexa,  I  have  studied  U.  S.  National  Museum 
cestode  No.  1110,  which  is  one  of  the  specimens  upon  which  Stiles 
(1896)  based  his  original  description  and  which  was  designated  by 
him  a  cotype.  Permission  was  given  to  dismount  and  section  the  speci- 
men. Inasmuch  as  it  had  been  mounted  in  balsam  for  18  years,  this 
was  obviously  a  difficult  task  to  accomplish;  but  by  careful  work  two 
series  of  sections  have  been  made,  which,  while  by  no  means  suited  for 
careful  study,  are  sufficient  to  show  that  all  the  differences  supposed 
to  exist  between  this  cestode  and  C.  mosaica  (Hall,  1908)  are  due  to 
errors  or  incorrect  conclusions  as  to  the  former.  These  points  will  be 
taken  up  one  at  a  time. 

The  cirrus  pouch  Stiles  (1896)  records  as  being  288  to  320/*  long 
in  C.  perplexa.  Eighteen  pouches  I  measured,  however,  were  all  530/* 
long.  Hall  found  the  pouch  in  C.  mosaica  to  be  475  to  640/*  long,  with 
an  average  length  of  550/*.  There  is  therefore  no  difference  in  this 
regard.  In  position  and  form  the  cirrus  pouch  and  vagina  are  iden- 
tical with  Hall's  descriptions.  The  specimen  at  hand  is  not  suited  for 
histological  study  and  comparison.  According  to  Stiles'  description 
the  testes  of  C.  perplexa  are  absent  from  the  median  part  of  the  pro- 


48  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [398 

glottid.  In  the  specimen  at  hand,  however,  they  extend  across  the  field 
continuously  in  a  manner  identical  with  the  description  by  Hall  for 
C.  mosaica.  There  is  of  course  no  reason  to  doubt  the  correctness  of 
Stiles '  observation ;  but  it  seems  evident  that  the  condition  he  described 
is  not  diagnostic.  As  mentioned,  I  have  found  the  testes  in  C.  pectinata 
almost  separated  in  two  fields. 

Hall  recognizes  in  his  specimens  some  differences  from  Stiles'  ac- 
count as  to  the  time  of  first  appearance  of  genital  organs.  He  acknowl- 
edges that  such  differences  may  be  due  to  differences  in  technique  and 
mentions  that  Stiles'  account  was  based  upon  unsectioned  material. 
I  have  not  attempted  to  section  the  anterior  end  of  the  specimen  at 
hand  since  it  is  not  in  condition  for  close  study.  However  there  is  no 
valid  reason  for  considering  the  specimens  different  on  this  point.  The 
mosaic  markings  mentioned  by  Hall  can  hardly  be  considered  of  im- 
portance; anyway,  they  are  shown  to  a  slight  extent  by  the  specimen 
at  hand. 

There  are  no  other  points  of  difference  between  the  two  accounts. 
I  have  compared  Stiles'  cotype  in  nearly  every  particular  with  Hall's 
account  and  find  only  two  points  of  difference:  (1)  eggs  appear  first 
in  the  uterus  in  the  94th  proglottid;  and  (2)  the  ovary  is  850/*  across, 
and  of  somewhat  different  appearance  from  the  condition  Hall  de- 
scribes. These  points  of  difference  are  taken  up  in  the  paragraph  after 
the  next. 

Specimens  of  this  form  from  cottontails  (Sylvilagus  floridanus) 
at  Blair,  Oklahoma,  show  the  ovary  exactly  as  described  by  Hall  and 
of  the  same  dimensions,  that  is,  about  600/*  across.  The  cirrus  pouch 
is  somewhat  smaller,  and  old  pouches,  when  gorged  with  sperm,  shorten 
up  and  become  pearshaped.  In  two  specimens  sexual  maturity  is 
reached  in  the  90th  proglottid.  In  other  points  they  do  not  differ  from 
Hall's  description.  Figures  46  to  49  are  from  this  material.  These 
Oklahoma  specimens,  it  will  be  seen,  resemble  the  Maryland  specimen 
in  that  maturity  is  reached  about  the  90th  proglottid.  The  gorging  and 
shortening  up  of  the  cirrus  pouch  suggests  an  explanation  of  the  basis 
of  Stiles'  measurements.  The  general  appearance  of  the  worms  is 
identical  with  both  the  Maryland  and  the  Colorado  forms,  and  the 
form  of  the  ovary  is  identical  with  that  of  those  from  Colorado.  It 
seems  very  probable  that  the  appearance  of  the  ovary  presented  by  the 
Maryland  specimen  is  due  to  the  flattening  process  to  which  it  was 
subjected  preparatory  to  mounting  in  toto.  The  lobes  stretching  out 
laterally  at  full  length  are  in  just  the  position  they  would  assume  with 
such  treatment.  As  to  the  difference  in  the  stage  at  which  maturity  is 
reached,  the  same  explanation  would  apply  here  as  for  Anoplocephala 


399]  ANOPLOCEPHALIDAi—DOUTHITT  49 

variabilis  (page  22).  Bowie,  Maryland,  and  Blair,  Oklahoma,  are  in  the 
Upper  Austral  faunal  zone,  while  Seven  Lakes,  Colorado,  is  in  the 
Boreal. 

The  Colorado  forms  may  deserve  the  rank  of  a  separate  variety, 
on  account  of  the  difference  in  stage  of  sexual  maturity;  but  until  the 
character  has  been  determined  for  more  than  two  or  three  individuals 
on  each  side,  such  a  step  would  not  be  justified.  Variations  of  far 
greater  import  have  been  recorded  in  this  paper  which  were  found  in 
individuals  of  the  same  species  and  from  the  same  host,  and  even  in 
different  proglottids  of  the  same  individual.  The  name,  Cittotaenia 
mosaica,  should  therefore  be  dropped,  and  the  excellent  description  by 
Hall  (1908)  applied  to  C.  perplexa,  the  only  change  necessary  being  to 
recognize  that  sexual  maturity  may  not  occur  until  the  95th  proglottid 
and  that  in  addition  to  occurring  in  Sylvilagus  pinetis  of  Colorado  it 
also  occurs  in  Sylvilagus  floridanus  of  Maryland  and  Oklahoma.  There 
seems  no  reason  to  doubt  that  the  testes  are  sometimes  separated  into 
two  groups,  as  described  by  Stiles. 

C.  pectinata  and  C.  perplexa  are  strikingly  similar  in  several 
unique  characters,  and  it  seems  not  improbable  that  their  differences 
may  be  due  to  environmental  conditions,  since  one  is  a  parasite  in  the 
cottontail  and  the  other  in  the  jack  rabbit;  but  my  present  opinion  is 
that  such  a  decision  would  be  premature,  without  testing  it  by  breeding 
experiments.  C.  perplexa  differs  from  C.  pectinata  in  the  following 
points,  all  of  which  are  quantitative: 

1.  The  testes  are  fewer 

2.  The  cirrus  pouch  is  smaller 

3.  There  is  a  more  complicated  network  of  excretory  ducts  in  the 
lateral  portions  of  the  median  field 

4.  The  ovary  is  smaller 

THE   GENUS   CITTOTAENIA 

Zschokke  in  1907  described  a  cestode  from  Phascolomys  wombat, 
giving  it  the  name  Moniezia  diaphana.  His  excellent  description  how- 
ever seems  to  be  that  of  a  perfectly  normal  Cittotaenia,  and  disagrees 
with  Moniezia  on  every  point  in  which  these  two  genera  differ.  The 
species  is  here  transferred  to  the  genus  Cittotaenia  for  the  following 
reasons : 

1.  The  vagina  is  not  ventral  on  the  right  and  dorsal  on  the  left, 
as  in  Moniezia. 

2.  There  are  no  shields  on  the  ends  of  the  horns  of  the  pyriform 
body. 


SO  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [400 

3.  The  uterus  is  tubular — not  reticular. 

4.  The  dorsal  excretory  duct  is  dorsolatral  to  the  ventral. 

Two  changes  should  be  made  in  the  diagnosis  of  the  genus  Citto- 
taenia.  It  should  be  stated  that  the  oviduct  joins  the  ovary  directly  in 
front  of  the  median  axis  of  the  vitelline  gland  and  that  the  uterus 
crosses  the  excretory  ducts  dorsally  only,  or  not  at  all. 

The  diagnosis  therefore  becomes  as  follows: 

Anoplocephalinae,  with  segments  broader  than  long.  Two  sets  of 
reproductive  organs  in  each  segment.  Genital  canals  pass  dorsal  of 
longitudinal  excretory  vessels  and  nerves.  Interproglottidal  glands 
absent.  Vagina  ventral  of  cirrus  pouch  on  both  sides  of  the  segment. 
Oviduct  joins  ovary  directly  in  front  of  the  median  axis  of  the  vitelline 
gland.  Uterus  either  confined  to  the  median  field  or  crossing  the  excre- 
tory ducts  dorsally  only.  Development  usually  by  simple  anterior  and 
posterior  outpocketing.  Eggs  with  well  developed  pyriform  apparatus 
of  ordinary  type,  or  without  pyriform  apparatus.  Adults  in  mammals 
and  birds. 

Type-species :    Cittotaenia  denticulata  Rudolphi  1804. 
The  genus  Cittotaenia  includes  the  following  species,  most  of  which 
are  fairly  well  known: 

1.  Marmotae-group.  Cirrus  pouch  pyriform,  distinct.  Dorsal 
duct  lateral  or  dorsal  of  ventral  (no  statement  concerning  C.  bursaria. 

C.  marmotae  Braun  C.  praecocquis  Stiles 

C.  denticulata  Rudolphi  C.  ctenoides  Railliet 

(C  latissima  Riehm  C.  diaphana  Zschokke 

(C.  goezei)  C.  zschokkei  von  Janicki 

C.  bursaria  von  Linstow  C.  quadrata  von  Linstow 

2.  Pectinata-group.  Cirrus  pouch  much  elongated,  resembling 
nozzle  of  a  hose,  and  indistinct;  becoming  pyriform  occasionally  in  old 
proglottids.    Dorsal  excretory  duct  median  to  ventral. 

C.  pectinata  Goeze  (in  part)  C.  rhea  Fuhrmann 
C.  variabilis  Stiles  C.  psittacea  Fuhrmann 

(C.  mosaica  Hall)  C.  avicolae  Fuhrmann 

C.  perplexa  Stiles  C.  kuvaria  Shipley 

COMPARATIVE  STUDIES  ON  THE  ANOPLOCEPHALIDAE 

Fuhrmann 's  classification  of  the  Cyclophyllidea  (1907)  which  is 
accepted  here  recognizes  in  the  subfamily  Anoplocephalinae  the  genera 
Triplotaenia,  Cittotaenia,  Moniezia,  Anoplocephala,  Andrya,  Bertiella, 
Schizotaenia  and  Aporna.     Each  of  these  genera  is  well  defined  and 


401]  AXOPLOCEPHALIDJE—DOUTHITT  51 

distinct;  and  except  for  Triplotaenia  and  Aporina  they  form  a  com- 
pact and  orderly  group.  But  little  attention  has  been  given  to  the 
relationships  of  these  genera  to  each  other.  In  the  following  pages  the 
various  resemblances  in  the  different  organs  are  considered,  with  refer- 
ence to  the  question  of  relationships  between  the  various  genera. 

1.  The  Structure  of  the  Uterus. — Two  general  types  of  uterus  have 
been  recognized  in  this  group :  the  tubular,  found  in  Bertiella,  Anoplo- 
cephala,  and  most  Cittotaeniae,and  the  reticular,  found  in  Andrya, 
most  Monieziae,  and  some  Cittotaeniae.  In  addition  the  uteri  of  Schizo- 
taenia,  Triplotaenia,  and  Aporina  have  been  recognized  as  each  being 
a  distinct  type  in  itself,  tho  the  uterus  of  Triplotaenia  is  not  really 
different  from  the  tubular.  The  uterus  of  Schizotaenia  has  been  shown 
in  this  paper  to  be  of  the  reticular  type,  the  reticulum  being  apparently 
in  the  process  of  degeneration.  There  appears  no  reason  to  regard  the 
uterus  of  Aporina  as  of  any  other  than  the  tubular  type.  As  to  the 
relationships  of  the  reticular  and  tubular  types  to  each  other,  very 
little  has  been  done  in  the  way  of  investigation  except  to  conjecture. 
Generally  the  reticular  type  has  been  supposed  to  be  derived  from  the 
tubular  by  the  outgrowth  and  anastomosis  of  proximal  and  distal 
branches.  Following  are  offered  the  conclusions  from  studies  of  the 
reticulate  uteri  of  different  species  of  Andrya,  made  with  the  view  of 
studying  the  relationships  of  these  two  types  to  each  other. 

In  the  earliest  stages  in  which  indications  of  the  uterus  were 
observed  it  occupies  the  entire  region  later  occupied  by  the  fully  devel- 
oped reticulum.  This  stage  is  long  before  any  lumina  appear.  The 
uterine  tissue  is  more  or  less  diffuse,  there  being  no  indication  of  segre- 
gation along  lines,  except  that  it  is  more  or  less  "stringy"  in  transverse 
axis.  As  the  proglottid  increases  in  size,  the  cells  segregate  more  and 
more  into  definite  lines  which  run  in  all  directions  and  connect  at  fre- 
quent intervals,  forming  a  network.  These  lines  of  segregation  do  not 
spread  from  any  central  locality  but  appear  in  situ  and  so  far  as  could 
be  observed,  at  about  the  same  time  in  all  parts  of  the  proglottid.  Later 
cavities  appear  in  these  tubes;  these  again  do  not  spread  out  from  any 
central  source  but  appear  in  situ  and  without  the  stimulus  of  entering 
ova.  When  fully  developed  they  form  a  continuous  network  of  open 
tubes.  As  these  tubes  become  filled  with  ova  they  expand  and 
coalesce,  forming  a  single  saccular  cavity.  Development  then  proceeds 
by  regular  outpocketing,  anteriorly  and  posteriorly.  Thus  in  the  early 
stages  of  development  there  is  no  indication  that  the  reticulate  uterus 
has  been  derived  from  a  transverse  tubular  type  by  outgrowth  and 
anastomosis  of  branches,  as  was  suggested  by  Stiles  (1896).  It  is  true 
of  course  that  the  uterus  spreads  from  some  original  source  in  the  course 


52  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [402 

of  development;  but  there  is  no  evidence  that  there  intervenes  a  trans- 
verse tubular  stage  which  is  comparable  in  any  sense  to  the  uterus  of 
Anoplocephala. 

On  the  other  hand  it  seems  much  more  probable  that  the  reticulate 
uterus  has  been  derived  by  lateral  spreading  from  a  median  longitu- 
dinal uterus,  such  as  we  find  in  more  primitive  cestodes,  and  is  the  first 
step  in  the  evolution  of  the  transverse  uterus  rather  than  the  last;  that 
the  tubular  uterus  of  Anoplocephala  has  been  the  product  of  simplifi- 
cation. Schizotaenia  seems  to  be  in  an  intermediate  condition ;  the 
uterus  appears  in  a  widely  diffused  condition  but  the  cavity  appears 
first  as  an  anterior  transverse  tube.  Some  such  view  becomes  necessary 
if  Andrya  is  regarded  as  a  primitive  type,  as  other  characters  indicate. 
It  is  a  matter  of  sincere  regret  to  me  that  other  matters  prevent  making 
a  careful  study  of  the  uterus  at  this  time. 

Aporina,  Andrya,  and  Anoplocephala  agree  in  that  the  uterus,  in 
nearly  all  species,  extends  laterad  beyond  the  excretory  ducts,  crossing 
them  ventrally.  Never  does  it  cross  them  dorsally  in  anlage  and  at  all 
stages  it  lies  mostly  ventral.  In  Schizotaenia,  and  apparently  always 
in  Bertiella,  it  is  confined  to  the  median  field  in  mature  proglottids 
and  in  its  fully  developed  stage  crosses  the  ducts  mostly  on  their  dorsal 
side,  if  at  all.  In  Cittotaenia  and  Moniezia  the  uterus  in  anlage  usually 
crosses  both  excretory  ducts  dorsally  but  in  some  species  is  confined  at 
all  stages  to  the  median  field.  Apparently  it  never  penetrates  between 
the  excretory  ducts  and  the  ventral  cortical  layer.  These  points  fur- 
nish very  important  indications  of  the  relationships  of  the  varied  genera 
to  each  other.  The  development  of  the  uterus  by  outpocketing  must 
be  recognized  as  a  very  general  character  of  the  subfamily,  and  in 
general,  a  primitive  one.  It  has  been  shown  to  be  the  method  of  devel- 
opment in  Andrya,  where  it  was  supposed  not  to  occur,  and  in  one 
species  of  Schizotaenia. 

2.  The  Distribution  of  Testes. — The  genus  Aporina  has  been  con- 
sidered unique  in  that  the  testes  extend  laterad  beyond  the  excretory 
ducts.  In  the  present  paper  however  are  described  four  species  of 
Andrya  and  two  of  Anoplocephala  in  which  the  testes  do  extend  across 
the  excretory  ducts  on  the  side  away  from  the  pore,  and  in  the  other 
species  of  Anoplocephala  here  taken  up  they  do  in  many  proglottids. 
Likewise,  Zschokke's  figures  for  Bertiella  edulis  (1899,  Taf.  XX,  Fig. 
2)  shows  that  in  this  species  the  testes  may  extend  nearly  across  the 
ducts.  It  seems  probable  that  they  occur  in  the  lateral  fields  in  all  or 
nearly  all  species  of  Anoplocephala  and  Andrya,  and  in  the  median 
field  in  all  species  of  Schizotaenia  and  most  species  at  least  of  Bertiella. 
The  double  pored  genera  of  course,  cannot  be  compared  in  this  regard. 


403]  ANOPLOCEPHALIDJE—DOUTHITT  53 

The  genus  Aporina  remains  distinct  in  that  the  testes  cross  the  excre- 
tory ducts  on  the  side  in  which  the  pore  would  be,  if  one  were  present. 
The  testes  of  Bertiella  and  most  species  of  Andrya  are  anterior; 
those  of  Andrya  occupy  almost  the  entire  median  field  except  for  the 
space  above  the  ovary.  Those  of  most  species  of  Schizotaenia  and  at 
least  some  species  of  Anoplocephala  are  mainly  posterior ;  and  for  Citto- 
taenia  and  Moniezia  they  are  posterior  in  all  cases  where  one  can  judge. 
However,  Andrya  translucida  has  the  testes  posterior  and  two  species 
of  Schizotaenia  have  them  mainly  anterior.  It  is  therefore  impossible 
to  form  any  final  conclusions  as  to  the  paths  of  evolution  in  this  regard. 
In  looking  however  for  the  type  that  has  the  most  generalized  distri- 
bution of  testes,  attention  is  at  once  attracted  by  the  conditions  found 
in  Andrya.  Here,  especially  in  Andrya  primordialis,  is  found  a  near 
approach  to  a  distribution  that  could  give  rise  by  suppression  to  any 
distribution  found  in  the  subfamily,  with  the  exception  of  Aporina. 

3.  The  Vaginal  Pore  and  Vagina. — The  position  of  the  vagina  and 
vaginal  pore  with  reference  to  the  cirrus  pouch  appears  from  these 
studies  to  be  the  most  stable  generic  character.  In  Aporina  and  An- 
drya the  vaginal  pore  and  vagina  are  directly  posterior  to  the  cirrus 
pouch,  which  position  is  primitive  among  cestodes.  In  Bertiella  the 
pore  and  vagina  are  either  posterior  or  somewhat  dorsal.  In  Anoplo- 
cephala the  pore  is  ventral  to  the  pouch,  and  the  vagina  is  ventral, 
and  for  at  least  the  lateral  half  of  its  length  is  not  posterior.  In 
Cittotaenia  the  pore  and  vagina  are  ventral  but  may  be  at  the  same 
time  posterior  also.  In  Moniezia  they  are  ventral  to  the  pouch  on  the 
right  side  and  dorsal  on  the  left  and  usually  they  are  also  somewhat 
posterior.  In  Schizotaenia  both  vagina  and  vaginal  pore  are  directly 
in  front  of  the  cirrus  pouch.  The  evidence  provided  here  points  to  the 
conclusion  that  the  three  genera  first  mentioned  are  the  most  primitive. 

4.  The  Ovary. — Two  types  of  ovary  can  be  distinguished.  In  one 
the  lobes  radiate  in  all  directions  in  the  horizontal  plane  from  the  point 
of  attachment  of  the  oviduct;  in  the  other  the  lobes  radiate  from  a 
transverse  bar,  which  forms  the  basal  part  of  the  ovary.  The  second 
type  is  perhaps  the  only  one  found  in  Anoplocephala  and  Schizotaenia, 
and  it  is  present  with  modifications  in  Moniezia  and  Cittotaenia.  The 
first  type  is  typically  represented  in  Andrya.  Unfortunately  most 
of  the  drawings  of  species  of  Bertiella  and  Aporina  are  obscure 
on  this  point.  At  any  rate  the  character  is  not  of  much  systematic 
importance  because  the  two  conditions  are  not  very  distinct  from  each 
other  and  are  obscured  by  modifications. 

5.  The  Excretory  ducts. — Nothing  is  known  as  to  the  excretory 
■ducts  of  Triplotaenia.     Bertiella  and  Moniezia  have  the  dorsal  ducts 


54  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [404 

either  dorsal  or  dorsomedian,  and  Cittotaenia  has  them  dorsal,  lateral 
or  dorsomedian.  Usually  the  dorsal  duct  is  lateral  in  Anoplocephala 
and  Schizotaenia ;  but  in  certain  stages  of  contraction  it  has  been  ob- 
served to  be  directly  dorsal  in  both  genera.  In  Andrya  the  dorsal  duct 
is  lateral  in  all  cases  observed.  In  the  scolex  of  all  species  of  Andrya 
and  Anoplocephala  which  were  observed  however,  the  dorsal  duct  is 
dorsal  while  in  Schizotaenia  anoplocephaloides  it  is  lateral  in  the  scolex. 
It  would  appear  therefore  that  little  evidence  as  to  relationship  is 
provided  by  the  excretory  ducts. 

THE  EVOLUTION  OF  THE  ANOPLOCEPHALIDAE 

The  following  arguments  favor  the  conclusion  that  Andrya  ap- 
proaches most  closely  of  all  the  genera  to  the  ancestral  type  of  the 
Anoplocephalidae. 

1.  In  Andrya  the  ovary  is  either  median,  or  very  close  to  the 
median  line,  and  none  of  the  female  glands  are  far  from  the  median 
line. 

2.  In  Andrya  the  vagina  and  vaginal  pore  are  directly  posterior 
to  the  cirrus  pouch. 

3.  In  Andrya  is  found  the  most  generalized  distribution  of  testes, 
with  the  possible  exception  of  Aporina. 

4.  In  the  generalized  species  of  Andrya  is  found  a  primitive,  un- 
modified vas  deferens,  which  character  is  found  in  no  other  genus  except 
Aporina. 

5.  In  Andrya  is  found  the  most  genuine  and  least  modified  reticu- 
late uterus. 

6.  In  Andrya,  the  uterus  in  anlage,  crosses  the  excretory  duct 
ventrally. 

As  characters  of  more  doubtful  value,  the  following  may  be  noted. 

1.  The  pyriform  body  is  simple  (no  pyriform  body  in  Aporina). 

2.  The  vesicula  seminalis  is,  in  the  most  generalized  species,  a 
simple,  globular  expansion. 

3.  The  genital  pores  are  in  some  species  irregularly  alternate. 
As  specialized  characters  of  Andrya  I  can  see  but  one ;  the  genital 

pores  are  either  dextral  or  tend  towards  dextrality,  whereas  one  would 
expect  the  ancestor  of  the  Anoplocephalidae  to  have  single,  irregularly 
alternating  genital  pores.  Such  a  point  constitutes  but  a  slight  depart- 
ure however  in  the  light  of  the  different  arrangements  that  have  arisen 
in  the  family;  and  in  this  regard  evolution  could  easily  reverse  its 
action.  The  possession  of  a  prostate  gland  would  seem  at  first  to  be  a 
specialized  character;  but  this  structure  appears  to  be  associated  with 


405]  AXOPLOCEPHALIDJE—DOUTHITT  55 

an  unmodified  vas  deferens  and  to  cease  to  have  a  purpose  for  existence 
when  the  vas  deferens  comes  to  function  as  a  storehouse  for  spermato- 
zoa, as  is  the  case  in  all  other  genera  except  Aporina. 

Within  the  genus  Andrya  there  has  been  a  wide  range  of  evolution. 
A.  primordialis  is  decidedly  the  most  primitive  in  structure  of  ovary 
and  uterus  and  testicular  distribution ;  and  only  in  the  arrangement  of 
genital  pores  is  it  unlike  what  should  be  expected  in  the  ancestors  of 
all  the  Anoplocephalidae.  In  Andrya  macrocephala  and  A.  translucida 
on  the  other  hand  is  found  a  rather  high  degree  of  specialization.  As 
regards  position  in  the  scale  Aporina  is  a  close  contestant  with  Andrya 
for  the  most  primitive  position,  each  having  distinct  points  in  its  favor. 
Aporina  must  be  regarded  however  as  an  offshoot  since  certain  charac- 
ters which  it  possesses  exclude  it  from  the  direct  line. 

From  forms  like  the  most  primitive  Andryae  there  sprang  appar- 
ently a  line  destined  to  give  rise  to  all  the  other  Anoplocephalidae, 
except  possibly  Bertiella.  The  characteristics  of  this  line,  many  of 
which  were  subsequently  modified,  were  as  follows: 

1.  The  lobes  of  the  ovary  did  not  radiate  from  the  point  of  attach- 
ment of  the  oviduct,  but  instead  extended  out  from  a  transverse  bar 
which  formed  the  base  of  the  ovary. 

2.  The  vagina  and  vaginal  pore  had  begun  to  move  anteriorly  and 
ventralwards. 

3.  The  testes  had  come  to  lie  mostly  in  the  posterior  part  of  the 
proglottid. 

4.  The  cirrus  pouch  had  begun  a  conspicuous  development  in  size, 
foreshadowed  in  Andrya  primordialis. 

5.  The  female  glands  remained  in  the  primitive  position,  near  the 
median  line. 

6.  The  prostate  gland  was  lost  and  the  vesicula  seminalis  was 
acquired. 

The  genus  Anoplocephala  apparently  separated  from  the  rest  early, 
and  developed  along  somewhat  different  lines.  It  retains  the  following 
Andryan  characters. 

1.  The  uterus,  in  anlage,  crosses  the  excretory  ducts  ventrally. 

2.  The  testes  extend  into  the  lateral  field  on  the  side  away  from 
the  pore. 

3.  The  proximal  end  of  the  oviduct  lies  beneath  the  median  lobe 
of  the  vitelline  gland. 

4.  The  tendency  towards  dextral  arrangement  of  genital  pores  is 
retained  and  perfected. 


56  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [406 

After  Anoplocephala  had  split  off  the  common  line  became  modified 
as  follows: 

1.  The  uterus  ceased  to  cross  the  excretory  ducts,  in  anlage. 

2.  The  ovary  moved  forward  and  laterad  so  as  to  lie  directly  in 
front  of  the  vitelline  gland  with  the  proximal  end  of  the  oviduct  in 
front  of  the  median  line  of  this  gland. 

3.  The  testes  become  confined  to  the  median  field. 

From  this  line  developed  Cittotaenia,  Schizotaenia,  and  Moniezia. 
Cittotaenia  resembles  Anoplocephala  in  some  respects  but  the  following 
evidences  of  relationship  to  Moniezia  outweigh  them: 

1.  In  "both,  the  uterus  crosses  the  excretory  ducts  dorsally  only, 
never  ventrally  as  in  Anoplocephala,  Andrya,  and  Aporina.  In  some 
cases  in  each  of  these  two  genera,  the  uterus  is  confined  to  the  median 
field  at  all  stages. 

2.  At  least  one  Cittotaenia  has  a  reticulate  uterus. 

3.  Both  have  two  sets  of  genital  organs. 

4.  The  dorsal  excretory  duct  is  dorsal  or  median  to  the  ventral 
in  most  species  of  each. 

The  points  arguing  for  close  relationship  between  Moniezia  and 
Schizotaenia  are  as  follows: 

1.  Each  retains  at  least  remnants  of  the  reticulate  uterus. 

2.  Some  individuals  of  Schizotaenia  show  asymmetry  in  regard  to 
the  relations  of  the  genital  glands. 

3.  Moniezia  has,  to  a  considerable  extent,  at  least,  the  mulberry- 
like  vitelline  gland,  such  as  is  found  in  most  species  of  Schizotaenia. 

4.  The  ripe  uterus  is  divided  into  compartments  in  some  species 
of  both  genera. 

Although  retaining  the  reticulate  uterus  Moniezia  and  Schizotaenia 
are  in  many  ways  the  most  specialized  representatives  of  their  subfam- 
ily, except  of  course  for  Triplotaenia.  They  have  specialized  along 
different  lines  and  the  decision  as  to  which  is  nearest  the  ancestral  line 
depends  upon  the  criteria  adopted. 

The  position  of  Bertiella  in  this  scheme  is  somewhat  doubtful.  I 
had  no  specimens  at  hand  for  study  and  so  must  depend  upon  the 
accounts  of  others.  The  following  points  argue  for  close  relationship 
with  the  higher  representatives  of  the  genus  Andrya  rather  than  with 
other  genera. 

1.  The  vagina  and  vaginal  pore  are  directly  posterior,  or  poste- 
rior and  dorsal  to  the  cirrus  pouch. 

2.  The  cirrus  pouch  is  either  identical  with  the  higher  Andryae 


407]  ANOPLOCEPHALIDJE—DOUTHITT  57 

in  form  and  size,  or  much  smaller  (the  pouch  is  larger  in  the  other 
genera). 

3.  The  testes  are  mainly  anterior. 

4.  In  species  concerning  which  information  is  available,  the  ovi- 
duct and  ovary  are  related  to  each  other  as  in  Andrya,  and  not  as  in 
Schizotaenia. 

The  following  points  argue  for  relationship  rather  to  Schizotaenia- 
like  forms,  however. 

1.  The  cirrus  is  sometimes  spiny. 

2.  The  genital  pores  are  sometimes  regularly  alternate. 

3.  The  uterus,  at  least  in  anlage,  is  confined  to  the  median  field. 

4.  The  dorsal  excretory  ducts  are  often  dorsomedian,  as  in  rela- 
tives of  Schizotaenia. 

5.  The  testes  seem  to  be  confined  to  the  median  field,  as  in  Schizo- 
taenia ;  in  some  species  of  Schizotaenia,  also,  the  testes  are  anterior. 

I  am  inclined  to  consider  the  aguments  for  closest  relationship  to 
the  genus  Andrya  as  the  most  important,  and  to  attribute  the  resem- 
blances to  Schizotaenia  to  parallel  evolution. 

The  relationships  of  the  genus  Triplotaenia  are  problematical  and 
will  probably  always  remain  uncertain.  The  possession  of  two  sets  of 
reproductive  organs  suggests  relationship  to  Cittotaenia  or  Moniezia. 
The  following  points  argue  for  relationship  to  Schizotaenia. 

1.  The  uterus,  in  anlage,  is  confined  to  the  median  field. 

2.  The  horns  of  the  pyriform  body  are  long  and  filamentous. 

3.  The  vagina  is  inconspicuous  and  apparently  ephemeral. 
These  points  argue  for  little  more  than  general  relationship  however 
and  all  of  them  might  easily  be  accidental. 

The  following  points  argue  that  Thysanosoma  is  most  nearly  related 
to  the  higher  representatives  of  the  Anoplocephalinae  and  is  descended 
from  forms  like  Moniezia  and  Schizotaenia. 

1.  There  are  sometimes  two  sets  of  reproductive  organs  in  Thy- 
sanomosa. 

2.  The  genital  ducts  sometimes  cross  the  excretory  ducts  ventrally 
in  Schizotaenia. 

3.  The  genital  organs  are  placed  far  laterad,  in  Thysanosoma. 

4.  The  vaginal  pore  may  be  dorsal,  or  anterior  to  the  cirrus  pouch, 
in  Thysanosoma. 

5.  Accessory  transverse  commissures  of  the  excretory  ducts  are 
present  in  species  of  both  Schizotaenia  and  Thysanosoma. 

These  arguments  however  are  at  best  inconclusive.  The  most  that 
can  be  said  is  that  Thysanosoma  seems  to  be  related  to  the  higher 
Anoplocephalinae . 


58  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [408 

That  Stilesia  and  Avitellina  are  quite  closely  related  to  Thysano- 
soma  is  too  evident  to  need  discussion.  The  exact  position  of  these 
genera  with  respect  to  each  other  must  be  left  unsettled,  however,  until 
the  species  of  Thysanosoma  have  received  more  careful  study.  At 
present  we  can  do  no  better  than  accept  Gough's  decision  (1911)  that 
Stilesia  and  Avitellina  represent  a  subfamily  distinct  from  Thysano- 
soma. 

In  the  subfamily  Linstowinae,  conditions  are  unsettled  and  uncer- 
tain, owing  to  the  recent  efforts  of  Beddard  (1911,  and  1912),  and 
others.  It  seems  little  short  of  absurd  to  decide  from  external  characters 
(1911:652)  as  to  which  of  about  20  genera  a  given  cestode  should  be 
referred  to,  the  possible  list  including  such  genera  as  Anoplocephala, 
Zschokkeella,  Thysanosoma,  Davainea,  Hymenolepis,  Oochoristica  and 
Taenia.  "Within  two  years,  Beddard  has  actually  placed  one  cestode 
(whose  latest  name  is  Zschokkeella  gambianum)  in  three  genera,  repre- 
senting two  subfamilies;  and  there  is  no  assurance  that  the  present 
disposal  is  final.  Since  Beddard  has  proved  his  worth  as  an  investigator 
by  his  researches  in  other  fields,  one  might  expect  better  things  in  the 
future ;  but  as  yet,  while  he  has  dwelt  at  great  length  upon  the  signifi- 
cance of  the  size  and  appearance  of  sucker  and  scolex,  the  presence  or 
absence  of  a  neck,  and  other  such  features,  he  has  failed  to  tell,  in  any 
instance,  whether  the  genital  ducts  cross  the  excretory  ducts  dorsally 
or  ventrally,  or  pass  between  them;  at  least,  several  hours  were  spent 
in  search  in  his  voluminous  articles  without  finding  any  information 
on  this  point.  Bischoff  (1912)  has  likewise  contributed  considerably 
towards  a  state  of  general  confusion  in  this  group.  He  has  added  to 
the  genus  Inermicapsifer  several  species,  without  giving,  apparently, 
any  adequate  study  to  each.  The  few  facts  that  he  gives  are  not  suffi- 
cient in  most  cases  to  show  that  the  species  exist ;  and  if  they  do  exist, 
there  is  no  evidence  in  many  cases  that  they  should  be  placed  in  the 
Inermicapsifer  group  rather  than  in  some  allied  group. 

In  the  present  condition  of  affairs,  there  are  no  characters  except 
trivial  unimportant  ones  upon  which  to  separate  the  so-called  genera 
Zschokkeella,  Inermicapsifer,  Thysanotaenia  and  Hyracotaenia  from 
each  other.  My  own  opinion  is  that  they  all  belong  in  one  genus,  with 
the  exceptions  of  Zschokkeella  remota  and  Thysanotaenia  lemuris  which 
seem  to  belong  elsewhere.  In  the  present  condition  of  our  knowledge 
of  most  of  the  species  of  the  group,  however,  any  alterations  would 
merely  add  to  the  confusion  already  existing.  "Taenia"  anoplocepha- 
loides  must  be  given  a  place  amongst  the  Linstowinae,  close  to  the  genus 
Linstowia.    Of  the  genera  of  the  Linstowinae,  Linstowia  shows,  by  the 


409]  AKOPLOCEPHALIDJE—DOUTHITT  59 

position  and  form  of  genital  organs  and  excretory  ducts,  that  it  is  the 
most  generalized,  unless  "Taenia"  anoplocephaloides  be  given  a  lower 
place.  The  following  facts  argue  that  Linstowia  is  descended  from  the 
higher  Anoplocephalinae,  from  forms  somewhat  like  Schizotaenia  and 
its  relative  Moniezia. 

1.  The  uterus,  in  some  Schizotaenia  and  Monieziae,  in  its  fully 
developed  stage  is  divided  up  into  compartments  much  like  the  embryo- 
capsules  of  the  Linstowinae. 

3.  The  uterus  of  the  Linstowinae,  in  anlage,  is  not  different  in 
type  from  the  diffuse  uterus  of  the  genus  Schizotaenia. 

3.  The  uterus  apparently  crosses  the  excretory  ducts  dorsally  in 
the  Linstowinae;  information  is  vague  or  wanting  as  to  most  species, 
however. 

4.  The  genital  ducts  have  been  observed  to  cross  the  excretory 
ducts  ventrally  in  some  individuals  in  Schizotaenia. 

5.  The  cirrus  pouch  is  unusually  large  in  Schizotaenia  and  Lin- 
stowia. 

The  following  diagram  represents  the  scheme  of  evolution  here 
proposed. 


AyvtelUna 

Thysanosoma 


\ntrtmcn)>iifer 
ZschcWcttllal     Thysanottnia 

StlUsia.  \        \  |    Triplatatnu 


? 
Schizotaenia 


Btrtulla 
5j)tciaUzed  Andrw  /         /  Aporinx 

Prinultvt  Andryat\ 

Ancestral   bnoflocephalui&e 


60  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [410 

LIFE  HISTORIES  OF  POCKET  GOPHER  ANOPLOCEPHALIDS 

Throughout  the  present  studies  attention  has  been  given  to  the 
problem  of  the  life-histories  of  the  cestodes  of  the  pocket  gopher.  This 
problem  has  been  attacked  in  three  ways:  by  experimental  feeding,  by 
examination  of  suspected  hosts  for  cysticercoids,  and  by  observations  on 
the  habits  of  the  gopher.  Either  the  first  or  second  method,  or  both, 
have  been  tried  out  on  all  the  parasites  and  commensals  of  the  gopher 
which  I  have  been  able  to  find,  with  only  negative  results  as  yet;  it  is 
my  intention  to  keep  steadily  at  this  quest  whenever  opportunity  offers. 
Observations  on  the  habits  of  the  gopher  however  have  yielded  infor- 
mation which  may  prove  of  decided  value  in  the  quest  of  the  intermedi- 
ate hosts  of  the  Anoplocephalidae,  and  this  information  is  given  here 
in  the  hope  that  the  facts  may  be  of  assistance  to  other  workers  who 
are  giving  attention  to  this  subject.  In  order  to  make  the  matter  clear 
a  brief  statement  is  necessary  concerning  the  habits  of  the  gopher. 

I  have  found  Geomys  personatus  always  in  infertile,  sandy  ground. 
Geomys  breviceps  I  have  caught  at  various  points  in  Texas  and  Okla- 
homa; but  nowhere  except  at  Norman,  Oklahoma,  have  I  caught  them 
in  anything  except  upland  regions  in  very  sandy  soil.  At  Norman, 
gophers  were  taken  along  a  stream  valley  in  soil  that  is  of  a  sandy  na- 
ture but  fertile  and  moist.  Geomys  bursarius  I  have  found  to  range 
over  about  every  conceivable  locality.  They  thrive  on  the  borders  of 
swamps,  on  hilltops,  on  open  plains,  in  heavy  pine  forests,  in  wind- 
blown sand,  and  in  heavy  clay  soil,  whether  uplands  or  lowlands. 

Wherever  he  occurs,  the  gopher  leads  a  life  wholly  apart  from  his 
fellows  except  at  the  mating  season.  His  tunnel  is  seldom  as  much  as 
50  meters  long.  This  tunnel  is  practically  the  gopher's  world  except 
during  the  mating  seasom  In  the  evening  he  may  come  to  the  top  of 
the  ground  for  a  few  moments;  but  he  does  not  wander  more  than  a 
few  feet  from  the  mouth  of  the  burrow.  In  regard  to  the  occurrence 
of  all  the  species  of  Anoplocephalidae  found  infesting  the  gopher,  one 
rule  has  proved  to  be  invariable :  they  occur  only  in  gophers  inhabiting 
rich  soils,  preferably  clay  soils,  but  occasionally  in  sand  when  it  is 
mixed  with  humus  and  is  of  a  swampy  nature.  To  this  rule  I  have 
found  not  a  single  exception.  Apparently  not  all  regions  of  fertile  soil 
are  infected,  and  lowlands  in  the  main  are  most  heavily  infected. 
Swampy  places  have  been  found  to  yield  the  best  of  all. 

A  specific  instance  is  illuminating.  At  Brainerd,  Minnesota,  fifteen 
gophers  were  caught  in  the  sandhills;  on  examination  one  cestode  was 
found  which  belonged  in  the  genus  Hymenolepis.  The  traps  were  then 
moved  to  Bean  Flat,  200  meters  away.     Bean  Flat  is  a  low,  almost 


411]  ANOPLOCEPHALIDJE—DOUTHITT  61 

swampy  region  in  which  the  gophers  can  live  only  along  the  margins 
of  the  low  river  terraces  which  cross  the  flat.  With  a  single  exception, 
every  adult  gopher  examined  was  infected,  this  locality  and  another  200 
meters  away  which  was  similar  yielding  five  species  of  Anoplocephalid 
cestodes  of  three  genera.  Twenty-six  specimens,  representing  three  spe- 
cies, were  taken  from  one  host.  When  one  considers  that  the  gopher 
never  wanders  more  than  a  few  feet  from  the  mouth  of  his  burrow 
except  during  the  mating  season  this  fact  becomes  all  the  more  signifi- 
cant. The  conclusion  is  forced  upon  the  student  that  there  is  some 
connection  with  the  nature  of  the  soil  that  determines  the  infection; 
it  seems  most  probable  that  the  intermediate  hosts  are  plant-feeding 
insects  that  live  only  in  fertile  soil  preferring  wet  lowlands. 

Since  this  rule  holds  true  for  six  species  of  four  genera  of  the 
Anoplocephalidae  it  seems  probable  that  it  will  hold  true  for  the  family 
generally;  that  there  is  something  associated  with  fertile  soils  and 
swampy  places  that  is  necessary  for  the  continued  existence  of  these 
parasites ;  that  if  cattle  and  sheep  could  be  confined  to  sandhill  pastures 
(which  assuredly  would  not  be  practical),  they  would  not  become  in- 
fected with  Thysanosoma  and  Moniezia.  This  rule  is  at  least  worth 
trying  out  by  other  workers  who  are  giving  attention  to  the  life-histories 
of  these  cestodes.  The  camel  cricket  (Ceuthophilus)  has  been  found  to 
abound  in  the  tunnels  of  the  pocket  gopher  in  localities  where  these 
cestodes  occur  and  to  be  absent  from  localities  where  the  cestodes  do 
not  occur.  However  all  attempts  to  infect  these  animals  have  resulted 
in  failures.  It  is  my  intention  to  continue  these  experiments  both  with 
these  animals  and  others  whenever  opportunity  offers. 

The  following  table  shows  the  infections  found  in  adult  gophers. 
All  gophers  that  were  less  than  8  months  old,  even  tho  fully  grown  were 
without  exception  uninfected.  The  separation  of  the  various  species 
of  Hymenolepis  which  were  found  was  rather  hastily  done  and  only 
approximate  correctness  was  aimed  at.  The  species  of  Cittotaenia  from 
Brainerd,  Minnesota,  was  unfortunately  immature  and  could  not  be 
worked  out.  It  is  not  C.  praecocquis  described  by  Stiles  (1896)  from 
the  same  host-species.  The  examinations  were  all  made  by  me,  with  the 
exception  of  those  from  Nebraska  for  which  I  have  to  thank  Dr.  John 
E.  Gutberlet. 


62 


ILLINOIS   BIOLOGICAL    MONOGRAPHS 


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413]  ANOPLOCEPHALIDAl—DOUTHITT  63 


KEY  TO  KNOWN  SPECIES  OF  ANOPLOCEPHALIDAE 

Owing  to  the  fact  that  the  literature  of  this  group  is  scattered, 
uncoordinated,  and  often  inaccessible,  the  task  of  identifying  material  is 
often  very  difficult  and  time-consuming.  The  writer  in  taking  up  for 
the  first  time  the  several  groups  here  studied,  has  had  good  cause  to 
realize  this.  In  the  hope  that  it  will  spare  to  others  this  waste  of  labor 
in  becoming  acquainted  with  the  field,  the  following  key  to  all  the 
known  species  of  Anoplocephalidae  is  presented.  A  great  many  poorly 
described  species  are  represented  here,  species  as  to  whose  affinities  no 
one  has  any  means  of  judging.  In  order  to  avoid  confusion  they  are 
left  in  the  genera  to  which  they  have  been  assigned,  tho  in  many  cases 
there  is  no  proof  that  they  belong  even  in  the  family.  To  prepare  a 
key  for  such  species  has  been  found  very  difficult  and  in  some  cases  the 
only  thing  that  could  be  done  was  to  list  them  with  such  descriptions 
as  are  possible. 

In  order  not  to  hamper  the  serviceableness  of  the  key  only  the  more 
important  synonyms  are  given  and  usually  only  the  best  reference  is 
given  for  each  species.  For  an  exhaustive  bibliographic  account  of  the 
various  species  and  genera,  the  reader  is  referred  to  Stiles  &  HassaU's 
excellent  summary  (1912).  True  phylogenetic  characters  have  been 
used  wherever  possible;  and  wherever  such  exist,  classifications  pre- 
viously proposed  have  been  accepted.  No  attempt  has  been  made  to 
treat  the  Linstowinae  critically ;  they  are  included  here  merely  to  make 
the  key  complete  and  critical  study  is  left  to  someone  better  fitted  to 
take  up  the  work. 

Family  Anoplocephalidae:  Scolex  and  suckers  unarmed.  Seg- 
ments nearly  always  broader  than  long,  generally  many 
times  broader  than  long.  Genital  pores  marginal  (absent 
in  Aporina).  Testes  numerous,  except  in  Triplotaenia. 
Uterus  transverse.  Embryos  with  thin,  transparent  shells, 
usually  with  a  pyriform  apparatus  borne  upon  the  inner- 
most shell.  Adults  in  mammals  and  birds,  mostly  in  her- 
bivorous mammals;  larval  stages  unknown.  Four  sub- 
families   - ~ 1 

1.  Genital  canals  dorsal  to  longitudinal  excretory  vessels  and 
nerves,  except  possibly  in  Triplotaenia,  and  in  some  pro- 
glottids  in  some  Schizotaenia.  Uterus  at  the  stage  of  sexual 
maturity  tubular,  netlike,  or  diffuse,  or,  in  Triplotaenia, 


64  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [414 

saclike.  Development  usually  by  outpocketing,  the  uterus 
rarely  breaking  up  into  separate  compartments,  never  with 
paruterine  organs.  Vitelline  and  shell  glands  always  pres- 
ent.   Anoplocephalinae    2 

Uterus  breaks  down  early  into  egg  capsules.  Genital  canals 
pass  ventral  to  excretory  ducts  or  between  them.  Cortical 
parenchyma  usually  very  thick.  A  single  set  of  reproduct- 
ive organs  in  each  segment  in  all  known  species.  Vitelline 
and  shell  glands  always  present — Linstowinae  51 

Paruterine  organs  present.  Genital  canals  pass  between  the 
excretory  vessels  and  dorsal  to  the  nerve.  Embryos  with 
two  shells,  usually  with  pyriform  apparatus  which  is  de- 
void of  horns.  Testes  in  two  lateral  groups  which  cross 
the  excretory  ducts  on  either  side.  Vitelline  and  shell 
glands  present Thysanosominae  70 

Paruterine  organs  present.  Genital  canals  pass  dorsal  to  excre- 
cretory  ducts  or  between  them.  Eggs  with  two  shells,  no 
pyriform  apparatus.  No  vitelline  gland  or  shell  gland. 
All  known  species  inhabit  ruminants.     (See  Gough,  1911.) 

Avitellininae  71 

Anoplocephalinae 

2.  Without  genital  pores.    Testes  crossing  excretory  ducts  on  both 

sides  of  the  proglottid.  No  pyriform  apparatus.  One  genus 

and  species.     (See  Fuhrmann,  1902.) Aporina  alba 

With  genital  pores.  Testes  not  crossing  excretory  ducts  on 
pore  side 3 

3.  With  not  more  than  one  cirrus  pouch  in  each  lateral  half  of  the 

segment.    With  numerous  testes.    With  external  evidences 

of  segmentation 4 

With  four  or  five  cirrus  pouches  in  each  lateral  half  of  the 
segment.  With  but  one  testis  in  each  lateral  half  of  the 
segment.  Without  external  evidences  of  segmentation.  One 
genus  and  species.     (See  von  Janicki,  1906.) 

Triplotaenia  mirabUis 

4.  With  but  one  set  of  reproductive  organs  in  each  proglottid 5 

With  two  sets  of  reproductive  organs  in  each  proglottid 6 

5.  Vaginal    pore    posterior   to    cirrus    pouch.     Uterus    reticular. 

Either  with  a  pedunculated  prostate  gland  or  with  ventral 
excretory  ducts  enormously  developed.  Genital  pores  all, 
or  nearly  all  dextral.     (See  present  paper.) —Andrya    7 


415]  ANOPLOCEPHALID.E—DOUTHITT  65 

Vaginal  pore  posterior,  or  posterior  and  dorsal  to  cirrus  pouch. 
Uterus  tubular.  No  prostate  gland.  Ventral  excretory 
ducts  normal  in  size.  Genital  pores  alternate,  except  in  the 
doubtful  Bertiella  pinguis  where  they  are  unilateral.  Tes- 
tes anterior.     (See  Bourquin,  1905.) Bertiella  12 

Vaginal  pore  and  vagina  ventral  to  cirrus  pouch.  Uterus  tubu- 
lar. No  prostate  gland.  Genital  pores  dextral  except  in 
the  doubtful  Anoplocephala  omphalodes  and  A.  minima. 
testes  only  on  the  side  away  from  the  genital  pore,  except  in 
A.  spatula  and  occasionally  in  A.  omphalodes.  (See  present 
paper.)    _ Anoplocephala  21 

Vagina  and  vaginal  pore  anterior  to  the  cirrus  pouch.  Uterus 
diffuse,  degenerate  reticular.  No  prostate  gland.  Testes 
either  posterior  or  anterior,  and  usually  mostly  on  the  pore 
side.     (See  present  paper.) _ Schizotaenia  30 

6.  Vagina  crosses  cirrus  pouch  ventrally  on  both  sides  of  the  seg- 

ment.   (See  Stiles,  1896,  and  present  paper.) Cittotaenia  34 

Vagina  crosses  cirrus  pouch  dorsally  on  the  left,  and  ventrally 
on  the  right  side.     (See  Stiles  &  Hassall,  1893.) Moniezia  44 

Genus  Andrya 

7.  Scolex  large.     Excretory  ducts  enormous.     No  prostate  gland. 

With  external  vesicula  seminalis „ 8 

Scolex  and  excretory  ducts  of  ordinary  dimensions.  With 
prostate  gland.  Vesicula  seminalis  confined  to  cirrus 
pouch _ 9 

8.  Length  10  to  20  cm.    Number  of  proglottids,  350  to  450.  Width 

of  strobila,  1.5  mm.    Vesicula  seminalis  very  large.    Host, 
Geomys  bursarius,  Minnesota.     (See  present  paper.) 

A.  macrocephala 
Length  9  to  12  cm.    Number  of  proglottids,  290.  Width  of  stro- 
bila, %  mm.     Vesicula  seminalis  only  two  or  three  times 
the  diameter  of  the  vas  deferens.    Host,  Geomys  bursarius, 
Minnesota.     (See  present  paper.) _ A.  translucida 

9.  European ;  over  60  cm.  long,  and  5  to  10  mm.  broad 10 

North  American ;  3  to  5  cm.  long,  not  over  2  mm.  wide _  11 

10.     Prostate  gland  elongate.     Testes  comparatively  few,  mostly  on 
the  side  away  from  the  pore.    Host,  Lepus  timidus,  Saxony. 

(See  Stiles,  1896.) A.  rhopalocephala 

Prostate  gland  round.  Testes  scattered  thru  entire  median 
field.  Hosts,  Lepus  cuniculus  and  L.  timidus,  Europe. 
(See  Stiles,  1896.)  A.  cunicuH 


66  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [416 

11.  Testes  transversely  elongated.     Genital   pore  near  middle  of 

lateral  margin.     Uterus  not  extending  ventral  to  ovary. 
Host,  Evotomys  gapperi  galei,  of  Colorado.     (See  present 

paper.)  _ — A.  communis 

Testes  not  transversely  elongated.  Genital  pores  near  distal 
end  of  lateral  margin.  Uterus  extends  ventral  to  ovary. 
Host,  Sciurus  hudsonica,  Minnesota.     (See  present  paper.) 

A.  primordialis 

Genus  Bertiella  (Synonym:  Bertia) 

12.  Genital  pores  alternate  13 

Genital    pores    unilateral.     Host,    Bucorax    abyssinicus.     Not 

properly  described,  position  not  assured.     (See  Fuhrmann, 
1904.)  _ B.  pinguis 

13.  Dorsal  excretory  duct  actually  dorsal.   Three  longitudinal  nerve 

trunks  on  either  side.    Cirrus  pouch  very  small  (except  in 

B.  poly orchis) .    Female  glands  in  a  dorsoventral  row 14 

Dorsal  excretory  duct  dorsomesial  to  ventral.  Cirrus  pouch  of 
ordinary  size,  and  well  developed  (except  in  B.  rigida). 
One  longitudinal  nerve  trunk  on  either  side  (no  statement 
for  B.  rigida).  Female  glands  arranged  in  a  transverse  row. 

Dorsal  excretory  duct  not  known,  apparently  absent.  One  nerve 
trunk.  Cirrus  pouch  of  ordinary  size,  egg-shaped.  Female 
glands  in  a  dorsoventral  row.  A  long,  convoluted  vas  def- 
erens. Host,  Lagidium  peruanum.  Not  properly  described. 
Assigned  by  von  Linstow  (1904)  to  the  genus  Bertia,  but 
it  is  by  no  means  certain  that  it  belongs  amongst  the  Ano- 
plocephalidae B.  forcipata 

The  following  poorly  known  species  is  placed  in  the  genus  Ber- 
tiella, perhaps  correctly  so.  Strobila  245  mm.  or  more  long 
by  10  mm.  broad.  Number  of  segments  about  350.  Genital 
pores  very  small,  irregularly  alternate.  Dorsal  canal  lateral 
to  ventral.  Cirrus  pouch  large  and  elongate.  Host,  Simia 
satyrus.     (See  Stiles,  1896;  or  R.  Blanchard,  1891.) 

B.  satyri 

14.  Genital  pores  regularly  alternate.    Receptaculum  seminis  poorly 

developed.     Ovary  extends  through  one-sixth  of  the  field 
between   the  excretory  ducts  and   is  not  bilobed.     Host, 

Troglodytes  niger,  Africa.     (See  Bourquin,  1905.) B.  studeri 

Genital  pores  alternate,  frequently  with  regularity.  Ovary  ex- 
tending through  about  one-fourth  the  width  of  the  proglot- 


417]  AX0PL0CEPHALIDA1—D0UTHITT  67 

tid,  divided  into  two  nearly  separated  lobes,  which  are 
connected  by  a  slender  neck.    Host,  Cercopithecus  callitri- 

chus.     (See  Beddard,  1911.) _ B.  cercopitheci 

Genital  pores  irregularly  alternate.    Ovary  not  bilobed „  15 

15.  Strobila  14  em.  or  more  long.    Receptaculum  seminis  globular. 

Eggs  enter  uterus  in  350th  segment.  Host,  Mycetes  niger, 
Paraguay.     (See  Meyner,  1895,  or  Stiles,  1896.) 

B.  mucronata 

Strobila  84  mm.  long.  Receptaculum  seminis  oval.  Eggs  enter 
uterus  in  130th  segment.  Host,  Macacus  radiatus.  (See 
Meyner,  1895,  and  Stiles,  1896.) „..B.  conferta 

Strobila  51  cm.  long.  Cirrus  pouch  of  ordinary  appearance. 
Host,  Macacus  cynomolgus,  India.  (See  von  Linstow, 
1 905. )   _ B.  poly  orchis 

16.  More  than  21  cm.  long.    Sexually  mature  proglottids  26  times 

as  broad  as  long,  ripe  proglottids  30  times  as  broad  as  long. 
Cortical  layer  exceptionally  thick,  in  young  proglottids 
forming  five-sixth  of  total  thickness,  in  older  proglottids 
about  three-fourths  of  thickness.  Female  glands  far  from 
median  line.  Testes  numbering  about  110.  Host,  Phalan- 
gista  sp.,  New  Guinea.     Not  adequately  described.     (See 

von  Janicki,  1905.)  B.  rigida 

Proglottids  not  so  short.    Cortical  layer  of  ordinary  thickness......  17 

17.  Cirrus  spiny — _ _  18 

Cirrus  not  spiny „  19 

18.  Length,  30  to  60  cm.    Number  of  proglottids,  600  to  850.    Eggs 

pass  into  uterus  in  300th  proglottid.  Testes  70  to  90.  Ovary 
extending  past  the  median  line  of  the  proglottid.  Host, 
Galeopithecus  volans,  Sumatra.     (See  Bourquin,  1905.) 

B.  elongata 
Length,  2  to  6  cm.  Number  of  proglottids,  80  to  200.  Genital 
pores  approach  regular  alternation.  Testes  50  to  70.  Ovary 
distant  by  more  than  twice  its  width  from  the  median  line. 
Host,  Galeopithicus  volans,  East  Indies,  Sumatra,  Java. 
(See  Bourqin,  1905.)  _ - B.  plastica 

19.  Vitelline  gland  posterior  to  the  ovary.    Length,  14  cm.    Num- 

ber of  testes  90,  extending  to  the  distal  end  of  the  proglot- 
tid. Diameter  of  ovary,  0.95  mm.,  not  greatly  displaced 
from  the  median  line,  its  mesial  edge  extending  across  the 
median  line.    Hosts,  doves  and  pigeons.     (See  Fuhrmann, 

1902.) B.  dclafondi 

Vitelline  gland  not  posterior  to  the  ovary 20 


68  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [418 

20.  Length,  50  to  60  cm.     Several  thousand  proglottids.     Testes 

about  200,  reaching  the  distal  limit  of  the  proglottid. 
Ovary  lying  one-third  beyond  the  median  line  of  the  pro- 
glottid. Host,  Phascolarctus  cinerus,  Australia.  (See 
Zschokke,  1898. )  B.  obesa 

Length,  66  cm.  Number  of  proglottids,  up  to  1500.  Testes 
fewer  than  in  B.  sarsinorum,  extending  nearly  to  the  pos- 
terior limit  of  the  proglottid.  Ovary  reaching  just  to  the 
median  line  of  the  proglottid.  Host,  Phalanger  ursinus, 
Australia  and  vicinity.     (See  Zschokke,  1899.)- B.  edulis 

Length,  7  cm.  Number  of  proglottids,  220.  Testes  30  to  40, 
mostly  in  a  single  row  at  the  anterior  end  of  the  proglottid. 
Ovary  distant  by  nearly  half  its  width  from  the  median 
line  of  the  proglottid.  Host, Phalanger  ursinus,  Australia 
and  vicinity.     (See  Zschokke,  1899.) B.  sarsinorum 

Genus  Anoplocephala  (Syn. :    Plagiotaenia) 

More  than  half  the  species  placed  in  this  genus  are  poorly 
known,  and  the  statements  made  are  often  not  dependable ; 
some  of  the  species  probably  do  not  exist,  and  others  prob- 
ably belong  even  in  other  families.  For  this  reason,  a  log- 
ical key  is  out  of  the  question,  and  no  attempt  has  been 
made  to  make  one;  the  following  arbitrary  division  is 
made. 

21.  At  least  fairly  well  known  species,  generic  position  assured „  22 

Improperly  described,  position  not  assured 27 

22.  Cirrus  pouch  enormous,  0.8  mm.  or  more  long,  lying  mostly 

within  the  excretory  ducts  23 

Cirrus  pouch  ordinarily  not  over  0.3  mm.  long,  more  than  half 
of  it  lying  outside  the  excretory  ducts 24 

23.  Length,  25  to  40  mm.    Number  of  proglottids,  30  to  40.    Sexual 

maturity  reached  in  10th  or  12th  proglottid.  Receptacu- 
lum  seminis  globular,  small.  Host,  Tapirus  americanus, 
Brazil.  Not  fully  known  anatomically;  all  points  known 
are  essentially  in  agreement  with  the  next  species.     (See 

Luhe,  1895.)  A.,  globiceps 

Length,  10  mm.  Number  of  proglottids,  10  to  28.  Sexual  matu- 
rity reached  in  eighth  or  ninth  proglottid.  Receptaculum 
seminis  large  and  oblong,  mostly  ventral  to  cirrus  pouch. 
Cirrus  not  spiny.  Ovary  not  bilobed.  Hosts,  Lepus  cunicu- 
lus  and  Lepus  variabilis,  France.     (See  present  paper.) 

A.  wimerosa 


419]  ANOPLOCEPHALIDJE—DOUTHITT  69 

Length,  6  to  30  mm.  35  to  55  proglottids.  Cirrus  spiny. 
Sexual  maturity  reached  about  the  15th  proglottid.  Host, 
Equus  caballus,  Europe.  (See  Stiles,  1896;  and  R.  Blanch- 
ard,  1891. )  „ A.  mamillana 

24.  Ovary  consists  of  two  practically  separate  lobes.    Receptaculum 

seminis    greatly    elongated.     Host,    Equus    caballus,    Old 

World.     (See  Kahane,  1880.)  A.  perfoliata 

Ovary  not  bilobed 25 

25.  (The  better  descriptions  are  not  accessible  to  me;  information 

compiled  from  several  sources).  9  to  80  cm.  long.  Scolex 
4  to  6  mm.  in  diameter.  Maximum  width,  5  to  20  mm. 
Synonys:    A.  plicata,  A.  zebrae,  etc.     (See  Scheibel,  1895.) 

A.  magna 
Scolex  considerably  less  than  1  mm.  broad.    Maximum  breadth 
not  over  5  mm „. 26 

26.  Over  3  cm.  long.    Over  125  proglottids.    Host,  Geomys  bursa- 

rius,  North  America.     (See  present  paper.) JL  variabilis 

Length,  12.5  to  20  mm.  Number  of  proglottids,  60  to  75.  Hosts, 
Geomys  bursarius  and  Microtus  sp.,  North  America.  (See 
present  paper. )  „ A.  infrequens 

27.  Genital  pores  irregularly  alternate.  Length,  12  to  21  cm.  Num- 

ber of  proglottids,  250  to  300.  Cirrus  spiny.  Hosts,  Mus 
arvalis  and  Mus  amphibius.  (See  von  Janicki,  1906;  and 
Stieda,  1862. )  — A.  omphalodes 

Genital  pores  regularly  alternate.  Length,  about  2  mm.; 
breadth,  about  0.34  mm.  Number  of  proglottids,  50  to  70. 
Host,  pheasant,  Italy.  Description  worthless;  nothing  to 
indicate  that  it  belongs  even  in  the  family,  if  it  exists.  (See 
Mello,  1912. )  „ A.  minima 

Genital  pores  unilateral „ 28 

28.  Testes  extending  across  median  field.     Length,  25  to  45  mm. 

Number  of  proglottids,  100.  In  Heterohyrax  mossambica. 
Most  likely  does  not  belong  in  this  genus.     (See  Bischoff, 

1912;  and  von  Janicki,  1910.) A.  spatula 

Testes  confined  to  the  side  away  from  the  genital  pore 29 

29.  Length,  16  cm.     Cirrus  spiny.    In  Arvicola  campestris.     (See 

von  Linstow,  1878;  and  von  Janicki,  1906.) A.,  inermis 

Length,  3  to  4  cm.    In  Agricola  campestris.    ( See  Moniez,  1891 ; 

and  von  Janicki,  1906.) _ A.  blanchardi  )/ 

Length,  about  8  mm.  Breadth,  5  mm.  Number  of  proglottids, 
40  to  50.  In  Arvicola  nivialis,  Italy.  Description  worth- 
less. Species  may  not  belong  to  family,  if  it  exists.  (See 
Galli-Valerio,  1905. )  - - A.  dentata 


70  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [420 

Length,  10  to  16  cm.  Breadth,  6  to  8  mm.  200  to  300  seg- 
ments. 60  to  80  testes.  In  Arctomys  sp.,  Turkestan.  (See 
Zschokke,  1888;  and  Stiles,  1896.) A.  transversaria 

No  information  available.  •  (See  Railliet,  1893.) A.  restricta 

Genus  Schizotaenia 

30.  Genital  pores  regularly  alternate.     Testes  posterior,  except  in 

occasional  proglottids  in  one  species.  Less  than  20  cm. 
long  _ m 31 

Genital  pores  irregularly  alternate.  Testes  posterior,  extending 
uniformly  across  the  median  field.  About  3  cm.  long.  Hosts, 
Dicotyles  albirostris  and  D.  torquatus,  Brazil.  Not  well 
known,  position  not  assured.    (See  Liihe,  1895.) 8.  decrescens 

Genital  pores  unilateral.  Size  enormous,  up  to  120  cm.  long, 
and  7.5  cm.  broad.    Testes  mostly  on  pore  side 33 

31.  Testes  extending  nearly  uniformly  across  the  median  field  in 

only  two  or  three  rows  anteroposteriorly.  Female  glands 
near  the  median  line  so  that  the  ovaries  of  the  two  sides 
overlap  for  most  of  their  width „ _.  32 

Testes  nearly  all  in  the  half  that  contains  the  genital  pore. 
Female  glands  far  from  the  median  line,  the  ovary  not 
reaching  the  median  line  of  the  proglottid.  Hosts,  Erethi- 
zon  dorsatus  and  E.  epixanthus,  North  America.  (See 
present  paper. )  „ S.  variabilis 

Testes  either  extending  across  the  median  field,  or  in  two 
groups  separated  from  each  other  by  the  vitelline  gland; 
in  either  case  mostly  on  the  side  away  from  the  genital 
pore,  where  they  are  in  six  or  seven  rows  in  the  antero- 
posterior axis.  Female  glands  quite  large  very  near  the 
median  line.  Host,  Oeomys  breviceps,  Oklahoma.  (See 
present  paper. )  _ S.  anoplocephaloides 

32.  Length,  14  cm.    Number  of  proglottids,  about  280.     Vitelline 

gland  compact,  formed  of  many  small  radiating  lobes. 
Walls  of  vesicula  seminalis  glandular  throughout.  Vaginal 
walls   glandular.     Host,    Hydrochaerns    capybara,   Brazil. 

(See  von  Janicki,  1906. )_ S.  hagmanni 

Length,  3  to  4  cm.  Number  of  proglottids,  about  100.  Vitelline 
gland  formed  of  a  large  median  and  a  smaller  lateral  lobe. 
Walls  of  vesicula  seminalis  not  glandular,  except  at  inner 
end.     Vagina    ephemeral,  walls     not    glandular.     Hosts, 


421]  ANOPLOCEPHALIDJE—DOUTHITT  71 

Erethizon  dorsatus  and  E.  epixanthus,  N.  America.  (See 
present  paper. )  S.  americana 

33.  Testes  relatively  few,  confined  to  the  anterior  portion  of  the 

proglottid,  mostly  on  the  pore  side.  Ovary  small,  pear- 
shaped.  Host,  Rhinoceros  sondiacns  (and  other  rhinocero- 
ses?)    (See  MacCallum  &  MacCallum,  1912;  Taenia  gigan- 

tea.)  8.  gigantea 

Testes  very  numerous,  filling  almost  the  entire  field  dorsally. 
Ovary  consists  of  a  transverse  portion  which  extends  nearly 
across  the  median  field,  and  of  numerous  clubshaped  lobes 
leading  from  it.  Host,  Rhinoceros  indicus  (and  other  rhi- 
noceroses?) (See  Deiner,  1912.)  For  reasons  for  trans- 
ferring to  Schizotaenia,  see  present  paper 8.  latissima 

Genus  Cittotaenia  (Syn. :  Ctenotaenia) 

34.  Cirrus  pouch  pyriform,  distinct.    Dorsal  excretory  duct  lateral 

or  dorsal  of  ventral  (no  statement  for  C.  bursaria) 35 

Cirrus  pouch  much  elongated  and  slender,  resembling  the  noz- 
zle of  a  hose,  and  less  distinct;  in  old  proglottids,  some- 
times becoming  pyriform  on  account  of  being  gorged  with 
sperm.    Dorsal  excretory  duct  generally  median 39 

35.  Uterus  not  a  simple  transverse  tube,  but  a  rather  simple  reticu- 

lum. Length,  40  to  80  cm.  Breadth,  15  mm.  Number  of 
proglottids,  200  or  more.  Genital  pores  in  posterior  corner 
of  segment.  Testes  very  numerous,  scattered  thru  the  dor- 
sal portion  of  the  median  field.     Host,  Lepus  cuniculus, 

Europe.     (See  Stiles,  1896,  and  others.) C.  denticulata 

Uterus  a  simple  transverse  tube 36 

36.  Testes  100  to  150,  confined  to  the  region  between  the  two  ova- 

ries, not  separated  into  two  lateral  groups.  Host,  Arcto- 
mys  sp.,  France.     (See  Blanchard,  1891;  and  Stiles,  1896.) 

C.  marmotae 

Testes  extending  nearly  or  quite  to  the  excretory  ducts  on 
either  side  anterior  to  the  ovaries,  and  not  separated  into 
two  lateral  groups 3T 

Testes  arranged  in  a  group  about  each  ovary,  absent  from  the 
median  part  of  the  field 38 

37.  Length,  55  mm.     Each  set  of  genital  organs  occupies  one-fifth 

the  cross  diameter  of  the  proglottid.  Cirrus  pouch  short 
and  clubshaped.  Testes  numerous,  spherical,  44yu.  in  diame- 
ter, reaching  the  posterior  end   of  the  proglottid  in  the 


72  ILLINOIS    BIOLOGICAL    MONOGRAPHS  1422 

median  part  of  the  field.  Host,  Lepus  nigricollis.  Local- 
ity, Nedunkeni.  Inadequately  described.  (See  von  Lin- 
stow,  1906.)  C.  bursaria 

Length,  180  mm.  Testes  very  numerous,  mostly  dorsal,  88/* 
long  by  53/t  broad.  Cirrus  spiny.  Each  ovary  occupies 
one-seventh  of  the  cross  diameter  of  the  proglottid.  Host, 
Lagidium  peruanum  cuvieri.  Locality,  Peru.  Not  well 
known.     (See  von  Linstow,  1904.) „ C.  quadrata 

Length,  40  mm.  Number  of  proglottids,  about  150.  Dorsal 
canal  lateral.  Host,  Oeomys  bursarius,  Iowa.  More  thor- 
ough description  needed.     (See  Stiles,  1896.) C.  praecocquis 

38.  Length,  80  cm.    500  to  750  proglottids.    60  to  80  testes  in  each 

group.  With  pyriform  body.  Hosts,  Lepus  cuniculus  and 
Lepus  cuniculus  domesticus.  Distribution,  Europe.  Not 
well  known.     (See  Stiles,  1896.) C.  ctenoides 

Length,  8  to  9  cm.  Number  of  proglottids,  250  to  350.  Testes 
about  15  or  20  on  each  side,  all  dorsal.  With  pyriform 
body.  (See  Zchokke,  1907,  under  name  Moniezia  diaphana; 
and  present  paper. ) C.  diaphana 

Length,  13  to  16  cm.  Three  longitudinal  excretory  ducts,  true 
dorsal  duct  outside.  Testes  30  on  each  side,  extending  thru 
medullary  portion.  No  pyriform  body.  (See  von  Janicki, 
1905.)   - C.  zschokkei 

39.  Diameter  of  scolex,  1.5  mm.    Cirrus  pouch  600/x  lond  and  20/x 

in  diameter.  Testes  about  110.  Host,  Rhea  americana,  of 
South  America.    Description  very  incomplete.     (See  Fuhr- 

mann,  1904. )  C.  rhea 

With  prostate  cells  on  vas  deferens _  40 

Without  prostate  cells  on  vas  deferens 41 

40.  Length,  10  cm.     Diameter  of  scolex,  230/i.     Testes  about  200, 

dorsal.  Cirrus  pouch  520/i  long  and  20/n  wide.  Host, 
Stringops  habroptUus.    Description  very  incomplete.     (See 

Fuhrmann,  1904.)  C.  psittacea 

Length,  15  to  22  cm.  Diameter  of  scolex,  800/x.  Testes  120  to 
140,  confined  to  the  region  between  the  ovaries.  Cirrus 
pouch  1  mm.  long,  120/x  in  diameter.  Host,  Anas  sp.  (See 
Fuhrmann,  1897.)  C.  avicolae 

41.  Dorsal  excretory  vessel  lateral  to  ventral.    Length,  5  cm.    Cir- 

rus pouch  300/*  long  and  40/i  broad.  Hosts,  Carpophaga 
vanwycki  and  Ptilopus  jambu.  Localities,  Kara  via  (New 
Britain)  and  Sumatra.  (See  Fuhrmann,  1902;  and  Ship- 
ley, 1900. )  ___ C.  kuvaria 


423]  ANOPLOCEPHALIDJE—DOUTHITT  ft 

Dorsal  excretory  duct  median  to  ventral „ 42 

42.  Testes  confined  to  the  region  between  the  ovaries.    Uterus  in 

anlage  crosses  excretory  ducts  dorsally.  Excretory  system 
simple.  Cirrus  pouch  400/x  long.  Hosts,  Sylvilagus  flori- 
danus  and  S.  palustris,  North  America.     (See  Stiles,  1896; 

Lyman,  1902;  and  present  paper.) C.  variabilis 

Testes  extending  laterad  beyond  the  ovaries.  Uterus  in  anlage 
confined  to  the  median  field.  Fifteen  or  more  large,  promi- 
nent ducts  crossing  the  proglottid  lengthwise  and  connect- 
ing the  transverse  commissures  of  the  excretory  ducts  with 
each  other _ 43 

43.  Cirrus  pouch  about  1  mm.  long,  extending  some  distance  mediad 

of  the  longitudinal  excretory  canals.  Testes  100  to  125. 
Hosts,  Lepus  timidus,  L.  variabilis,  and  L.  calif ornicus, 
Europe  and  North  America.     (See  Stiles,  1896;  Lyman, 

1902;  present  paper,  and  others.) C.  pectinata 

Cirrus  pouch  not  over  640/u.  long,  often  much  shorter.  Testes 
60  to  80.  Hosts,  Sylvilagus  floridanus  and  8.  pinetis.  (See 
Hall,  1908,  under  name  C.  mosaica;  and  present  paper.) 

C.  perplexa 


Genus  Moniezia  (Syn.  partim:  Paronia) 

44.  Description    not    accessible. — Moniezia    beauforti   von    Janicki 

1906b.    Host,  Cyclopsittacus  dioptahlmus,  in  Sekanto. 

M.  beauforti 

Length,  20-27  cm.  Host,  Macropus  giganteus,  Australia.  Poorly 
known,  position  uncertain.  (See  R.  Blanchard,  1891;  or 
Dujardin,  1845 :593.)  - M.  f estiva 

Uterus  tubular  or  slightly  branched,  not  reticular,  confined  to 
median  field  in  anlage.  In  its  final  stages  saccular,  or  with 
anterior  and  posterior  outpocketings.  Embryos  without 
pyriform  apparatus.  Dorsal  excretory  duct  dorsal  to  ven- 
tral (not  known  in  M.  variabilis).. 45 

Uterus  a  very  complicated,  finely  meshed  reticulum,  usually ,if 
not  always,  crossing  the  excretory  ducts  dorsally  in  anlage. 
Embryos  usually  with  pyriform  apparatus,  the  horns  of 
which  generally  end  in  a  disk.  Dorsal  excretory  duct  me- 
dian to  ventral _ - _ 47 

45.  Uterus  in  anlage  inverted  U-shaped,  embracing  ovary „  46 

Uterus  in  anlage  a  transverse  bar,  with  a  few  simple  branches 

at  its  median  end.     Communication  of  ripe  uteri  of  two 


74  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [424 

sides  of  proglottid  slight.  Number  of  testes  about  200. 
With  prostate  cells  on  vas  deferens.  Cirrus  pouch  90/x 
long.  Host,  PtUopus  sp.,  Sumatra.    (See  Fuhrmann,  1902.) 

M.  columbae 

46.  Uteri  of  two  sides  never  united.    Testes  about  140.    No  prostate 

cells  on  vas  deferens.  Cirrus  pouch  300  to  450/i  long. 
Hosts,  Ptilonopus  sp.,  Lorius  erythrothorax,  Trichoglossus 
novaehoUandiae,  Cyclopsittacus  suavissimus,  and  other  par- 
rots, New  Guinea  and  Sumatra.     (See  Fuhrmann,  1902; 

also,  Diamare,  1900  and  1901.) M.  carrinoi 

Uteri  of  two  sides  becoming  connected  very  early,  anteriorly. 
Number  of  testes,  about  100.     Host,  Chrysotis  amazonica, 

South  America.     (See  Fuhrmann,  1902.) M.  ambigua 

Uteri  of  two  sides  becoming  connected  distally.    Hosts,  4  species  of 
Rhamphastos.     Description  poor.     (See  Fuhrmann,  1904.) 

M.  variabilis 

47.  With  interproglottidal  glands,  which  are  linear,  and  are  not 

grouped  around  blind  sacs  (very  indistinct  in  M.  bene- 
deni.)    Planissima-growp 48 

With  interproglottidal  glands,  which  are  grouped  around  blind 
sacs.    Expansa-group 49 

No  interproglottidal  glands.    AZ&a-group 50 

48.  Length,  1  to  2  m.    Ripe  segments  12  to  26  mm.  broad,  and  1 

to  1.75  mm.  long.  Interproglottidal  glands  large,  distinct. 
Hosts,  Bos  taurus  and  Ovis  aries.  (See  Stiles  &  Hassall, 
and  others.) — _ M.  pianissimo 

Length,  4  m.  Ripe  segments  12  mm.  broad  by  3  mm.  long. 
Interproglottidal  glands  indistinct  (absent?).  Hosts,  Bos 
taurus  and  Ovis-aries.  Better  study  needed.  (See  Stiles 
&  Hassall,  1893.)  M.  benedeni 

About  0.66  m.  long.  Largest  segments  8  mm.  broad  by  1.5 
mm.  long.  Testes  in  a  quadrangle.  Interproglottidal 
glands  small.  Host,  Ovis  aries.  Better  study  needed.  (See 
Stiles  &  Hassall,  1893.) _ M.  neumanni 

49.  Length,  4  to  5  m.     Testes  in  mature  proglottids  usually  in  a 

quadrangle,  rarely  in  two  triangles.  Hosts,  Bos  taurus 
and  Ovis  aries.    See  Stiles  and  Hassall,  1893 ;  Tower,  1900 ; 

present  paper,  and  others.) M.  expansa 

Length,  about  1  m.  Testes  less  numerous  and  smaller  than 
in  M.  expansa.  Host,  Coassus  sp.,  Trinidad,  South  Amer- 
ica.   Poorly  known.     (See  Stiles  and  Hassall,  1893.) 

M.  oblongiceps 


425]  ANOPLOCEPHALIDAi—DOUTHITT  75 

Length,  1.6  to  2  m.  Testes  usually  in  two  triangles,  absent 
from  median  part  of  the  field.  Orifices  of  suckers  slitlike. 
Host,  Ovis  aries.     (See  Stiles  and  Hassall,  1893.) 

M.  trigonophora 

50.  Length,  0.6  to  2.5  m.     Testes  in  a  quadrangle.     Genital  pores 

in  anterior  half  of  the  segment.  Hosts,  Bos  taurus  and 
Ovis  aries.  Description  very  incomplete.  (See  Stiles  & 
Hassall,  1893.)  M.  alba 

Length,  41  cm.  Testes  in  entire  median  field,  except  part  occu- 
pied by  female  glands.  No  disk  figured  on  pyriform  body. 
Host,  Hippopotamus  amphibius.  Description  poor,  generic 
position  not  established.     (See  von  Linstow,  1901.) 

M.  amphibia 

Length  probably  not  over  10  cm.  Poorly  described.  Host, 
Ateles  hypoxanthus,  Brazil.    See  Liihe,  1895) M.  rugosa 

Linstowinae 

51.  Genital  pores  alternate.    Cirrus  pouch  well  developed.    Female 

glands  near  the  median  line 52 

Genital  pores  unilateral.  Cirrus  pouch  weakly  developed.  Fe- 
male glands  far  from  the  median  line,  except  Thysanotaenia 
lemuris  and  Zschokkeella  remota 53 

52.  Genital  canals  ventral  to  excretory  canals  and  nerve.    Cortical 

layers  of  great  thickness Linstowia,  54 

Genital   canals  between    excretory   canals.     Cortical   layer   of 

ordinary  thickness  (See  Fuhrmann,  1902.) 

"Taenia"  anoplocephaloides 

53.  For  the  following  so-called  genera,  no  differences  of  generic  im- 

port are  known,  with  the  possible  exceptions  of  Thysano- 
taenia lemuris  and  Zschokkeella  remota.  It  seems  most 
probable  that  they  all  belong  in  the  genus  Zschokkeella, 
with  the  two  exceptions  mentioned.  Most  of  the  species  are 
poorly  described,  so  no  attempts  at  rearrangement  have  been 
made.  They  are  listed  here  merely  to  make  the  key  com- 
plete. 

Genus  Zschokkeella 57 

Genus  Inermicapisfer „  58 

Genus  Thysanotaenia 68 

Genus  Hyracotaenia  _ 69 


76  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [426 

Genus  Linstowia 

54.  Cirrus  pouch  very  large,  reaching  to  the  median  line  of  the 

proglottid  or  nearly  so 55 

Cirrus  pouch  relatively  small,  not  reaching  to  the  lateral  excre- 
tory trunks.  Embryos,  in  ripe  proglottids,  mostly  in  lateral 
portions  of  the  segment 56 

55.  Testes  about  40,  mostly  posterior.     Host,   Perameles  obesula, 

Australia.    (See  Zschokke,  1898.) L.  semoni 

Testes  about  100,  mostly  anterior.  Host,  Echidna  hystrix,  Aus- 
tralia. (See  Zschokke,  1898.)  Tidswell,  1910,  has  declared 
Taenia  echidnae  Thompson  1893  and  Taenia  phoptica  Cob- 
bold  1879,  to  be  synonyms.  ( I  have  been  unable  to  find  this 
article,  or  even  to  learn  its  title) L  echidnae 

56.  Number  of  proglottids,  160  to  200.    Genital  pore  but  slightly  in 

front  of  the  middle  of  the  proglottid.  Ovary  of  about  8 
slender  lobes,  four  of  which  extend  towards  each  lateral  mar- 
gin.    Host,  Peramys  americana,  South  Americana.     (See 

Zschokke,  1904.) L.  jheringi 

Number  of  proglottids,  90.  Genital  pore  very  near  the  anterior 
end  of  the  margin.  Ovary  consists  of  two  comparatively 
compact  lobes  each  with  very  short,  slender,  radiating  pro- 
cesses.    Host,  Didelphys  tristriata,  Brazil.     (See  von  Jan- 

icki,  1906.) JL.  brasilensis 

Genus  Zschokkeella  (Syn.  Zschokkea  Fuhrman  1902) 

57.  Excretory  vessels  far  medial.    Testes  about  140,  largely  outside 

the  median  field.  Host,  Numida  ptilorhyncha...  (See  Fuhr- 
mann,  1902.) _ Z.  linstowi 

Testes  in  median  field.  Cirrus  pouch  round,  small,  one- 
eighteenth  the  diameter  of  the  proglottid.  Very  poorly 
described.  Most  of  the  facts  known  point  to  the  conclusion 
that  this  cestode  does  not  belong  in  the  genus  Zschokkeella ; 
but  not  enough  is  known  to  justify  placing  it  elsewhere. 
Host,  Oprecopithecus  pyrrhonotus,  West  Africa.  (See  von 
Linstow,  1905. ) „ Z.  remota 

Excretory  ducts  lateral  in  position.  Testes  in  two  groups, 
mostly  on  the  side  away  from  the  pore.  Host,  Cricetomus 
gambianum.     (See  Beddard,  1911,  1912.) Z.  gambianum 


427]  ANOPLOCEPHALIDJE—DOUTHITT  77 

Genus  Inermicapslper 

58.  No  information  of  importance.    (See  von  Janicki,  1910.)  _ 

/.  gondokorensis 

Testes  filling  most  of  median  field _ 59 

Testes  posterior,  extending  across  field 60 

Testes  in  two  lateral  groups „ 64 

59.  70  to  80  embryo-capsules,  with  8  to  10  embryos  in  each.    Num- 

ber of  proglottids,  70  to  80.    Length  of  cirrus  pouch,  400/*. 

Host,  Hyrax  sp.     (See  von  Janicki,  1910.) J.  pagensteckeri 

30  to  50  embryo  capsules,  up  to  6  embryos  in  each.  Number  of 
proglottids,  40.  Length  of  cirrus  pouch,  270/*.  Host, 
Hyrax  sp.    Locality,  Schoa.     (See  Bischoff,  1912.) L  paronae 

60.  Number  of  embryo  capsules,  100  or  more.    Number  of  proglot- 

tids, 300  or  more 61 

Number  of  embryo  capsules,  75  or  less.  Number  of  proglottids 
130  or  less.... _  62 

61.  (No  valid  point  of  distinction  from  next.     Length,  111  mm. 

Number  of  proglottids,  300.     Host,  Hyrax  capensis.     (See 

von  Janicki,  1910. ) „ /.  critica 

Length,  up  to  350  mm.  Number  of  proglottids,  400  to  500. 
Host,  Procavia  sp.,  South  Africa.     (See  von  Janicki,  1910.) 

I.  hyracis 

62.  Number  of  embryo  capsules,  75 ;  5  to  7  embryos  in  each.    Num- 

ber  of   proglottids,    130.     Number   of   testes,    80.     Host, 

Procavia  sp.,  South  Africa.     (See  von  Janicki,  1910.) 

7.  interposittis 
Number  of  embryo  capsules,  24  to  28.    Less  than  60  testes 63 

63.  Length,  40  mm.    8  to  10  embryos  in  each  capsule.    50  to  60  testes. 

Ho8t,Hyrax,  sp.,  Erethrea.     (See  Bischoff,  1912.) 

I.  abyssinicus 
Length,  15  to  20  mm.    5  embryos  in  each  capsules.    40  testes. 

Host,  Hyrax  sp.,  East  Africa.     (See  Bischoff,  1912.) „ 

7.  prionodes 

64.  Number  of  proglottids,  200.     Host,  Procavia  capensis.     (See 

Beddard,   1912. ) J.   capensis 

Number  of  proglottids,  not  over  70 65 

65.  5  to  7  embryos  in  each  capsule 66 

10  to  15  embryos  in  each  capsule 67 

66.  Cirrus  pouch  130  /t  long.    Host,  Hyrax  sp.,  East  Africa.     (See 

Bischoff,  1912. ) _ „.. J.  parvulxis 


78  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [428 

Cirrus  pouch  270  /x  long.  Host,  Hyrax  sp.,  East  Africa.  (See 
Bischoff ,   1912. ) _ I.   apospasmation 

67.  20  to  70  proglottids.     Host,  Procavia  sp.,  South  Africa.     (See 

von  Janicki,  1910.) 1.  settii 

12  to  15  proglottids.  Host,  Hyrax  sp.,  Rikwasee.  (See  Bisch- 
off, 1912.) _ 1.  lopas 

Genus  Thysanotaenia 

68.  Female  glands  near  the  median  line  of  the  proglottid.    No  dor- 

sal excretory  duct.  Host,  Lemur  macaco.  (See  Beddard, 
1911b,   1912. ) T.  lemuris 

Genus  Hyracotaenia 

69.  Ovary  entirely,  or  partly  laterad  of  ventral  excretory  duct. 

Vitelline  gland  dorsal.  Host,  Procavia  capensis.  (See  Bed- 
dard, 1912. ) H .  procaviae 

Ovary  entirely  mediad  of  excretory  ducts.  Vitelline  gland  ven- 
tral.   Host,  Procavia  capensis.     (See  Beddard,  1912.) 

H.  hyracis 

Thysanosominae 

Genus  Thysanosoma 

70.  Length,  15  to  30  cm.    Two  sets  of  reproductive  organs.    Testes 

in  median  field.  Posterior  flap  of  segments  fimbriate.  With 
pyriform  body,  which  has  no  horns.  Reported  from  several 
species  of  Cervus,  Cariacus,  Bos  &  Ovis.  (See  Stiles  & 
Hassall,  1893. )  T.  actinioides 

Length,  1  to  2  m.  Seldom  more  than  one  set  of  reproductive 
organs  to  the  proglottid,  pores  alternating  irregularly. 
Testes  in  lateral 'fields.  Posterior  flap  of  segments  fimbri- 
ate. With  pyriform  apparatus  which  has  no  horns.  Hosts, 
Ovis,  Bos,  and  Sus.     (See  Stiles  &  Hassall,  1893.) T.  giardi 

Length,  1.5  m.  Strobilization  not  distinct.  Genital  pores  alter- 
nating rather  regularly.  No  pyriform  apparatus.  Host, 
Capreolus  pygardus,  Siberia.  Description  unsatisfactory. 
(See  Kholodskovi,  1902.) T.  pygardi 

Avitellininae 

71.  Uterus  single ;  a  single  paruterine  organ.    Testes  in  four  groups. 

Genital  canals  pass  dorsally  of  both  excretory  ducts 

Avitellina  72 
Uterus  double;  two  paruterine  organs.     Testes  in  two  groups. 
Genital  canals  pass  between  excretory  canals Stilesia  73 


429]  ANOPLOCEPHALIDAE— DOUTHITT  79 

Genus  Avitellina 

72.  Length,  2  or  3  meters.     Host,  Ovis  aries,  Africa,  Italy.     (See 

Gough,  1911. ) A.  centripunctata 

Genus  Stilesia 

73.  Testes  mostly  median  or  dorsal  to  the  ventral  canal.     Hosts, 

ruminants,  Africa.     (See  Gough,  1911.) S.  hepatica 

Testes  all  lateral  to  the  ventral  canal 74 

74.  Vas  deferens  forms  a  dense  packet  of  convolutions  (functionally 

a  vesicula  seminalis)  between  nerve  and  ventral  canal  be- 
fore reaching  cirrus  pouch.     Host,  Qamelus  dromedarius, 

India  and  Algiers.     (See  Gough,  1911.) S.  vittata 

Vas  deferens  forms  at  most  3  or  4  loose  convolutions  between 
the  nerve  and  ventral  canal,  before  reaching  the  cirrus 
pouch.  Hosts,  Ovis  &  Capra,  Europe  and  India.  (See 
Gough,  1911. ) _ .#.  globipuncia 

SUMMARY  AND  CONCLUSIONS 

1.  The  cestodes  of  the  subfamily  Anoplocephalinae  are  in  some 
way  dependant  upon  rich  soils  for  their  existence  and  they  thrive  best 
in  wet  lowlands.  The  evidence  points  to  the  conclusion  that  the  inter- 
mediate hosts  are  some  group  of  insects  which  is  confined  to  such  regions ; 
and  since  the  hosts  of  the  Anoplocephalidae  are  almost  exclusively 
herbivorous,  it  would  seem  as  if  this  host  were  a  small,  plant-feeding 
insect. 

2.  The  primitive  Anoplocephaline  uterus  was  of  the  reticulate  type, 
which  in  turn  was  derived  from  a  median  longitudinal  tubular  uterus  by 
lateral  outgrowths.  The  transverse  tubular  and  diffuse  uteri  of  this 
group  have  been  derived  from  the  reticular  by  simplification. 

3.  In  the  early  primitive  Anoplocephalidae  the  uterus  crossed  the 
excretory  ducts  ventrally;  subsequently  it  became  restricted  to  the 
median  field  and  later  came  to  cross  the  excretory  ducts  dorsally. 

4.  The  position  of  the  vaginal  pore  and  vagina  is  one  of  the  most 
stable  anatomical  characters  of  the  Anoplocephalinae  and  should  be 
given  recognition  as  one  of  the  most  important  criteria  of  relationship. 
The  primitive  position  of  the  vagina  was  posterior  to  the  cirrus  pouch. 

5.  The  more  generalized  representatives  of  the  genus  Andrya  ap- 
proach the  nearest  of  all  known  Anoplocephalidae  to  the  ancestral  types 
of  the  family.    Leaving  out  of  consideration  the  aberrant  Triplotaenia, 


80  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [430 

the  genera  Moniezia  and  Schizotaenia  constitute  the  highest  types  of  the 
Anoplocephalinae ;  the  other  subfamilies  of  the  family  seem  to  have 
sprung  from  forms  like  these  two  genera. 

6.  "Cestodes  exhibit  a  high  range  of  variability  which  should  be 
taken  strictly  into  account  in  morphological,  phylogenetic,  and  syste- 
matic studies.  Generalizations  not  founded  upon  plenty  of  material  and 
careful  study  and  reflection  do  more  harm  than  good.  The  following 
reference  list  of  variations  and  anomalies  reported  in  this  paper  shows 
the  need  of  extreme  caution  in  coming  to  conclusions  regarding  struc- 
ture and  consequent  relationships. 

Andrya  communis — relation  of  cirrus  pouch  to  excretory 

ducts - page     9 

translucida — genital  functions - _ „..page  14 

excretory  ducts _ page  15 

Anoplocephala  variabilis — variations  due  to  state  of  contraction  ...page  23 

variations  due  to  habitat page  24 

Schizotaenia  variabilis — size  and  distribution  of  testes page  33 

excretory  system  page  34 

anoplocephaloides — course  of  genital  ducts „ page  36 

position  of  cirrus  pouch „ page  36 

distribution  of  testes __ page  37 

form  of  ovary __ page  38 

excretory  ducts page  39 

Cittotaenia  pectinata — distribution  of  testes... _ page  46 

number  of  proglottids — . page  46 

perplexa — distribution  of  testes page  47 

size  and  shape  of  cirrus  pouch page  47 

perplexa  and  pectinata  compared page  48 


\ 


431]  ANOPLOCEPHALIDAi—DOUTHITT  81 


BIBLIOGRAPHY 
Beddard,  F.  E. 

191  i.    Contributions  to  the   Anatomy  and   Systematic  Arrangement  of  the 

Cestoidea.    I.  On  Some  Mammalian  Cestoidea.     Proc.  Zool.  Soc,  London, 

191 1 .626-660. 
1911b.    Contributions  to  the  Anatomy  and  Systematic  Arrangement  of  the 

Cestoidea.    II.  On  Two  New  Genera  of  Cestodes  from  Mammals.     Proc. 

Zool.  Soc,  London,  1911:994-1018. 
1912.    Contributions   to   the   Anatomy  and   Systematic  Arrangement  of  the 

Cestoidea.    IV.  On  a  Species  of  Inermicapsifer  from  the  Hyrax,  and  on 

the  Genera   Zschokkeella,   Thysanotaenia  and   Hyracotaenia.     Proc.   Zool. 

Soc,  London,  1912:576-007. 

Bischoff,  C.  R. 

1912.    Cestoden  aus  Hyrax.     Zool.  Anz.,  39:751-758. 

Blanchard,  R. 

1 891.  Notices  helminthologiques  (deuxieme  series).  Mem.  soc.  zool  France, 
4:420-489;    38    figs. 

BOURQUIN,  J. 

1005.  Cestodes  de  Mammiferes.  Le  Genre  Bertia.  Revue  Suisse  Zool., 
13:415-506;  pi.  7-9- 

COHN,  L. 

1906.  Zur  Anatomie  zweier  Cestoden.  Centralblatt  f.  Bakt.,  I,  Abt,  Orig., 
40 :362-367. 

Deiner,  E. 

1912.  Anatomie  der  Anoplocephala  latissima  (nom.  nov.).  Arb.  Zool.  Inst. 
Wien,  19:347-372. 

DlAMARE,  V. 

1900.  Paronia  carrinoi,  n.  g.,  n.  sp.  von  Tanioiden  mit  doppelten  Geschlects- 
organen.  Centralbl.  f.  Bakt.,  1  Abt.,  28:846-851.  Also  in  Boll.  Mus.  Z00L, 
Genoa,  no.  91   (in  Italian). 

1901.  Zur  Kenntniss  der  Vogelcestoden.  Centralbl.  f.  Bakt,  1.  Abt.,  30: 
369-373. 

DUJARDIN,  F. 

1845.    Historie  naturelle  des  helminthes  ou  vers  intestinaux.     Paris,  1845. 

FUHRMANN,  O. 

1897.  Sur  un  nouveau  tenia  d'oiseau  (Cittotaenia  avicolae).  Revue  Suisse 
Zool.,  5:107-117;  pi.  5. 

1902.  Die  Anoplocephaliden  der  Vogel.  Centralbl.  f.  Bakt.,  1,  Abt.,  Orig., 
32:122-147. 

1904.    Neue  Anoplocephaliden  der  V6gel.     Zool.  Anz.,  27:384-387. 

1907.  Die  Systematik  der  Ordnung  der  Cyclophyllidien.  Zool.  Anz.,  32: 
289-297. 

1908.  Die  Cestoden  der  Vogel.     Zool.  Jahrb.,  Supp.,  10:1-232. 


82  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [432 

Galli-Valerio,  B. 

1905.    Einige   Parasiten  von   Arvicola   nivialis.     Zool.   Anz.,  28:519-522. 
Gough,  L.  H. 

191 1.  A  Monograph  of  the  Tapeworms  of  the  Subfamily  Avitellininae,  being 
a  Revision  of  the  Genus  Stilesia,  and  an  Account  of  the  Histology  of 
Avitellina  centripunctata  (Riv).  Quart.  Journ.  Micr.  Sci.,  56:317-385; 
pi.   12-14. 

Hall,  M.  C. 

1908.  A  New  Rabbit  Cestode,  Cittotaenia  mosaica.  Proc.  U.  S.  Nat.  Mus., 
34:691-699. 

1912.  The  Parasite  Fauna  of  Colorado.    Col.  Coll.  Pub.,  Sci.  Ser.,  12,  no.  10. 
Janicki,  C.  von. 

1905.  Beutlercestoden  der  Niederlandischen  Neuguinea — Expedition.  Zu- 
gleich  einiges  Neue  aus  dem  Geschlechtsleben  der  Cestoden.  Zool.  Anz., 
29:127-    . 

1906.  Studien  an  Saiigetiercestoden.  Zeitschr.  f .  wiss.  Zool.,  81 :505-595 ; 
pi.  20-25. 

1906b.    Die  Cestoden  Neu-guineas.    Nova  Guinea  Leiden,  5:181-200. 
1910.    Die  Cestoden  aus  Procavia.     Denkschriften  med.-naturw.  Ges.   Jena, 
16:373-396. 
Kahane,  Z. 

1880.     Anatomie  von   Taenia   perfoliata,   Goeze,   als   Beitrag  zur  Anatomie 
der  Cestoden.    Zeitschr.  f.  wiss.  Zool.,  34:175-254;  pi.  8. 
Kholodskovi,  N.  A. 

1902.    Contributions   a   la   connaisance    des    tenias    des    ruminants.     Arch. 
Parasit.,  6:145-148;  pi.  1. 
Leidy,  J. 

1855.    Notices  on  some  Tapeworms.     Proc.  Acad.  Nat.  Sci.,  Phila.,  7:433. 
Linstow,  O.  VON. 

1878.    Neue  Beobachtungen   an  Helminthen.    Arch.   f.  Naturges.,  1 :2i8-245 ; 

pis.  7-9- 
1 901.     Helminthen  von  den  Ufern  des  Nyassa-Sees, — ein  Beitrag  zur  Hel- 
minthen-Fauna   von    Sud-Afrika.     Jen.    Zeitschr.    f .    Naturw.,  35 :409-428 ; 
pis.  13-14. 

1904.  Helminthologische  Beobachtungen.  Centralbl.  f.  Bakt,  1.  Abt.,  Orig. ; 
37 :678-683. 

1905.  Neue  Helminthen.    Archiv  f .  Naturg.,  71,  1  -.267-276 ;  pi.  10. 

1905b.  Helminthen  aus  Ceylon  und  Arktischen  Breiten.  Zeitschr.  f.  wiss. 
Zool.,  82:187-193;  pi.  13.      1 

1906.  Helminthes  from  the  Collection  of  the  Colombo  Museum.  Spolia 
Zeylanica,  Colombo,  Pt.  II,  III,  pp.  163-188,  with  plates. 

Luhe,  M. 

1895.  Mitteilungen  iiber  wenig  bekannte  bez.  neue  sudamericanische  Tae- 
nien  des  k.  k.  naturhistorischen  Hof-Museums  in  Wien.  Archiv  f.  Na- 
turg., 61,  1:199-212;  pi.  11. 


433]  AN0PL0CEPHAL1DAZ—D0UTH1TT  83 

Lyman,  R.  A. 

1902.    Studies  on  the  Genus  Cittotaenia.     Trans.  Amer.  Micr.  Soc,  23:173- 
190;  pis.  26-27. 
MacCallum,  G.  A.  and  MacCallum,  W.  G. 

1912.    On   the    Structure    of   Taenia   Gigantea.     Zoolog.   Jahrb.,    Syst,   32: 

379-384. 
Mello,  U. 

1912.    Anoplocephala   minima,   n.   sp.    del    fagiano.     Monit.   Zool.    ItaJ,.,   23: 
124-130. 
Meyner,  R. 

1895.      Zwei    neue    Taenien    aus    Affen.      Ein    Beitrag    zur    Kenntniss    der 

Cestoden.     Zeitschr.   f.  Naturwiss.,  68:1-106;   pi.  2. 
Reprinted  exactly  except  for  title  as 

1895.     Anatomie  und  Histologic  zweier  neuen  Taenien,  Arten  des  Subgenus 
Bertia.     Taenia  (Bertia)   mucronata  n.  sp.  und  Taenia   (Bertia)   conferta, 
n.  sp.  Dissertation,  Halle. 
Moniez,  R. 

1891.    Notes  sur  les  Helminthes.     Revue  biol.  >Nord  France,  4:22-34,  65-79, 
108- 1 18. 
Parona,  Corrado. 

1900.     Helminthen  ex  Conradi  Paronae  Museo  Catalogus  (Sect.  2.  Cestodes). 
Genova,  October,  1900. 
Railliet,  A. 

1893.    Traite  de  Zoologie  medicale  et  agricole.    2.  ed.    Paris.    1893. 
Ransom,  B.  H. 

1909.    Taenioid  Cestodes  of  North  American  Birds.     Bull.  U.  S.  Nat.  Mus., 
69,  141  PP. 

SCHEIBEL,  A. 

1895.    Der    Bau    der    Taenia    magna    Abildgaard    (T.    plicata    Zeder),    ein 
Beitrag  zur  Kenntnis  der  Pferdetanien.     Dissertation.     Giessen. 
Shipley,  A.  E. 

1900.    A   Description  of   the  Entozoa  Collected  by   Dr.   Willey  During  his 
Sojourn  in  the  Western  Pacific. 

A.  Willey's  Zoological  Results,  Part  5,  pp.  552-556.    Fig.  23-26. 
Stieda,  Ludwig. 

1862.     Ein    Beitrag   zur   Kenntniss   der   Tanien.     Archiv    f.    Naturg.,   28,    I  : 
200-209. 
Stiles,  C.  W. 

1895.  Notes   on    Parasites. — 38.     Preliminary   Note   to   "A   revision   of   the 
adult  leporine  cestodes".    Vet.  Mag.,  2:341-346. 

1896.  A  Revision  of  the  Adult  Tapeworms  of  Hares  and  Rabbits.     Proc. 
U.  S.  Nat.  Mus.,  19:145-235;  pi.  5-25. 


~J 


84  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [434 

Stiles,  C.  W.  &  Hassall,  A. 

1893.     A  Revision  of  the  Adult  Cestodes  of  Cattle,   Sheep,  and  allied  ani- 
mals.   Bur.  An.  Ind.  Bull.  4. 
1902-1912.    Index  Catalogue  of  Medical  and  Veterinary  Zoology.     Authors' 

Index.    Bur.  An.  Ind.  Bull.  39. 
1912.     Index-Catalogue    of    Medical    and    Veterinary    Zoology.     Subjects : 
Cestoda  and  Cestodaria.     Hyg.  Lab.  Bull.  85. 
Tower,  W.  L. 

1900.    The    Nervous    System    of    the    Cestode    Moniezia    expansa.     Zool. 
Jahrb.,Anat.,  13:359-384;  pis.  21-26. 
Zschokke,  Fritz. 

1898.  Die   Cestoden   der   Marsupialia  und   Monotremata.     Denkschr.   med.- 
naturw.  Ges.  Jena,  8:358-380;  pi.  24. 

1899.  Neue  Studien  an  Cestoden  aplacentaler  Saugethiere.     Zeitschr.  f.  wiss. 
Zool.,  65:404-446;  pis.  20,  ax. 

1904.    Die    Darfncestoden    der    americanischen    Beuteltiere.      Centralblatt    f. 

B'akt.,  1.  Abt.,  Orig.,  36:51-621  pi.  1. 
1907.     Moniezia   diaphana,    n.   sp.     En   weiterer   Beitrag   zur   Kenntnis   der 

Cestoden  aplacentaler  Saugetiere.     Centralbl.  f.  Bakt.,  1.  Abt.,  Orig.,  44: 

261-264. 


435] 


ANOPLOCEPHALIDAZ—DOUTHITT 


85 


EXPLANATION  OF  PLATES 

Unless  otherwise  stated  all  drawings  were  drawn  to  scale,  measure- 
ments being  made  with  an  ocular  micrometer,  and  are  dorsal  side  upper- 
most, with  the  anterior  end  towards  the  top  of  the  plate. 


ABBREVIATIONS   USED 


c 

genital  cloaca 

RS 

CP 

cirrus  pouch 

SG 

ExD 

dorsal  excretory  vessel 

T 

ExV 

ventral  excretory  vessel 

UD 

ExT 

excretory  tubes  connecting  trans- 

Ut 

verse  commissures. 

Va 

N 

longitudinal  nerve  trunk 

VC 

0 

ovary 

VD 

Ovd 

oviduct 

VD 

Par 

parenchyma-filled      spaces      be- 

VDef 

tween    the    compartments    of 

VE 

the  uterus 

VG 

PG 

prostate  gland 

VS 

receptaculum  seminis 

shell  "gland 

testis 

uterine  duct 

uterus 

vagina 

ventral  transverse  commissure 

vas  deferents  (Plates  V  and  VI) 

vitelline  duct  (Plates  I  and  III) 

vas  deferens  (Plate  I) 

vasa  efferentia 

vitalline  gland 

vesicula  seminalis 


> 


86  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [436 


EXPLANATION  OF  PLATE 

Figures  i  to  4.    Andrya  primordialis. 

Fig.  1.     Proglottid  at  sexual  maturity,  dorsal  view;  ovary  and  uterus  omitted. 

Fig.  2.     Proglottid  at  sexual  maturity,  ventral  view,  to  show  ovary  and  uterus. 

Fig.  3.     Uterus  of  nearly  ripe  proglottid. 

Fig.  4.     Scolex,  drawn  from  temporary  mount. 
Figures  5  to  8.    Andrya  communis. 

Fig.  5.     Proglottid  at  sexual  maturity,  dorsal  view;  ovary  and  uterus  omitted. 

Fig.  6.     Proglottid  at  sexual  maturity,  ventral  view,  to  show  ovary  and  uterus. 

Fig.  7.    Ventral  view  of  vitelline  and  shell  glands  and   female  genital  ducts. 

Fig.  8.    Diagram  of  excretory  ducts  and  cirrus  pouch,  showing  a  different  ar- 
rangement from  that  shown  in  Figure  5. 
Figures  9  to  II.    Andrya  macrocephala  (see  also  next  plate). 

Fig.    9.  Mature  proglottid,  dorsal  view,  uterus  omitted. 

Fig.  10.  Mature  proglottid,  ventral  view,  to  show  uterus. 

Fig.  11.  Fully  developed  uterine  embryo,  middle  membrane  omitted. 


PLATE   I 


437]  AN0PL0CEPHAUDJE—D0VTH1T7  & 


PLATE  II 


88  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [438 


EXPLANATION  OF  PLATE 

Figures  12  and  13.    Andrya  macrocephala  (see  also  Plate  I). 

Fig.  12.    Uterus  of  ripe  proglottid. 

Fig.  13.    Section  of  scolex. 
Figures  14  to  16.    Andrya  translucida. 

Fig.  14.     Mature  proglottid,  dorsal  view. 

Fig.  15.    Mature  proglottid,  dorsal  view ;  ovary  and  shell  gland  omitted. 

Fig.  16.    Mature  proglottid,  ventral  view,  showing  ovary  and  uterus. 
Fig.  17.    Anoplocephala  wimerosa.    Mature  proglottid,  dorsal  view. 
Figures  18  to  21.    Anoplocephala  variabilis  (see  also  Plate  III). 

Fig.  18.    Section  of  scolex  in  expanded  condition. 

Fig.  19.     Section  of  scolex  in  contracted  condition. 

Fig.  20.    Uterus  of  ripe  proglottid. 

Fig.  21.    Uterine  embryo,  middle  membrane  not  represented. 


mf^Vr^fTXv^ 


y     f     >r     «f    "»> 


t JL_E_JF-    |9 


PLATE  II 


439]  AN0PL0CEPHALWAZ-D0UTH1TT  89 


PLATE  III 


90  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [440 


EXPLANATION  OF  PLATE 

Figures  22  to  24.    Anoplocephala  variabilis  (see  also  Plate  II). 

Fig.  22.    Mature  proglottid^  in  contracted  condition;  some  structures  omitted. 

Fig.  23.    Mature  proglottid  in  expanded  condition. 

Fig.  24.    Vitelline  gland,  shell  gland,  and   female  ducts ;  expanded  condition, 
ventral   view. 
Figures  25  to  27.     Anoplocephala  infrequent. 

Fig.  25.  Mature  proglottids  in  ventral  view,  some  structures  omitted.  The 
transverse  comissures  of  the  ventral  ducts  which  appear  straight 
here  are  sinuous  in   dorsoventral   plane. 

Fig.  26.     Section  through  the  scolex. 

Fig.  27.    Vitelline  gland,  shell  gland,  and  female  ducts;  ventral  view. 


PLATE  III 


441]  ANOPLOCEPHALIDJE—DOUTHITT  91 


PLATE  IV 


92  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [442 


EXPLANATION  OF  PLATE 

Figures  28  and  29.    Schizotaenia  americana. 

Fig.  28.    Mature  proglottid,  dorsal  view;  uterus  not  included. 

Fig.  29.  Mature  proglottid,  showing  uterus  (dotted),  oviduct,  and  shell  gland. 
Figures  30  to  32.    Schizotaenia  variabilis. 

Fig.  30.  Mature  proglottid,  dorsal  view.  After  this  drawing  was  completed 
the  study  of  additional  material  showed  that  it  is  not  normal  in  some  respects, 
namely:  (i)-the  testes  are  usually  further  forward,  anterior  to  the  transverse 
commissure;  (2)  the  accessory  ventral  commissures  were  found  in  only  one 
specimen;  (3)  the  vesicula  seminalis  is  usually  not  thrown  into  wide  curves  as 
here  shown.  The  uterus  may  be  somewhat  more  extensive  than  here  shown;  the 
poor  condition  of  the  material  does  not  permit  exact  judgment  as  to  the  limits  of 
this  structure. 

Fig.  31.  Diagram  of  excretory  system  of  five  adjacent  proglottids,  showing 
accessory  transverse  commissures  (AVC)  and  other  features.  The  absence  of 
the  main  transverse  commissure  in  some  cases  may  be  due  to  the  fact  that  the 
proglottids  are  not  fully  mature. 

Fig.  32.  Drawing  showing  the  cloaca  only  partly  everted  and  the  cirrus  pouch 
lying  consequently  mostly  mediad  of  the  excretory  ducts. 


PLATF.  IV 


443]  ANOPLOCEPHALIDJE—DOUTHITT  93 


PLATE  V 


94  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [444 


EXPLANATION  OF  PLATE 

Figures  33  to  40.    Schisotaenia  anoplocephaloides. 

Fig.  33.     Proglottid  just  before  sexual  maturity,  dorsal  view ;  uterus  and  vas 
deferens  omitted. 

Fig.  34.     Proglottid  at  sexual  maturity.     Ovary  and  testes  omitted.     Uterus 
dotted. 

Fig.  35.    Drawing  to  show  the  testes  extending  across  the  median  field,  the 
usual  condition  in  the  species. 

Fig.  36.    Ripe  proglottid,  showing  fully  developed  uterus  with  short  anterior 
and  posterior  outpocketings. 

Fig.  37.     Section  of  scolex  thru  suckers. 

Fig.  38.    Scolex  and  anterior  regions  from  toto  mount 

Fig.  39.    Fully  developed  uterine  embryo,  middle  embryonic  membrane  omit- 
ted. 

.  Fig.  4a    Diagram  to  show  excretory  system  in  a  series  of  proglottids.    Drawn 
ventral  side  uppermost. 


PLATE  V 


445]  ANOPLOCEPHALIDJE—DOUTHITT  95 


PLATE  VI 


96  ILLINOIS    BIOLOGICAL    MONOGRAPHS  [446 


EXPLANATION  OF  PLATE 

Figures  41  to  43.    Moniesia  expansa. 

Fig.  41.  Mature  proglottid,  dorsal  view;  shell  gland  and  part  of  testes  and 
genital   ducts   omitted   from   right  side  of  proglottid. 

Fig.  42.  Uterus  as  it  appears  just  prior  to  the  entrance  of  the  eggs,  a  compli- 
cated network  of  interconnecting  small  tubes. 

Fig.  43.  Uterus  in  its  final  stages,  a  system  of  separated  or  nearly  separated 
compartments. 

Figure   44.     Cittotaenia  variabilis.     Proglottid   at   sexual   maturity,   dorsal   view ; 
parts  of  genital  ducts  and  female  glands  omitted  from  left  side. 
Figure  45.     Cittotaenia  pectinata  americana.     Proglottid  at  sexual  maturity,  dor- 
sal view. 
Figures  46  to  49.  Cittotaenia  perplexa  from  Sylvilagus  floridanus  alacer,  Oklahoma. 

Fig.  46.  One-half  of  proglottid  at  sexual  maturity.  The  cirrus  pouch  is  here 
shown  to  reach  across  both  excretory  ducts:  usually  it  reaches  just  to  the  lateral 
nerve  trunk.  In  other  specimens  at  hand  the  testes  are  somewhat  more  numerous 
in  the  median  part  of  the  field,  and  less  numerous  in  the  lateral  parts.  In  Stiles' 
figures  they  are  wholly  absent  from  the  median  part  of  the  field. 

Figures  47  to  49.  Cirrus, pouches,  showing  variations  in  size  and  form.  All 
drawn  to  the  same  scale. 


PLATE  VT 


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UNIVERSITY  OF  ILLINOIS-URBANA 

rO.SILL  C004 

ILLINOIS  BIOLOGICAL  MONOGRAPHS  URBANA 
1  1914-15 


0112 


017753242 


